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Madagascar 

An environmental 
profile 



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Compiled by 

IUCN Conservation Monitoring Centre 



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MADAGASCAR 
AN ENVIRONMENTAL PROFILE 




WORLD CONSERVATION MONITORING CENTRE 
219 Huntingdon Road, Cambridge CB3 ODL U.K. 



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UNEP 



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INTERNATIONAL UNION FOR CONSERVATION OF NATURE 
AND NATURAL RESOURCES 

UNITED NATIONS ENVIRONMENT PROGRAMME 



MADAGASCAR 
AN ENVIRONMENTAL PROFILE 



Edited by M.D. Jenkins 



Prepared by 

The IUCN Conservation Monitoring Centre 

Cambridge, U.K. 



Published by IUCN, Gland, Switzerland and Cambridge, U.K. 
A contribution to GEMS - The Global Environment Monitoring System 

1987 



Prepared and published by IUCN, Gland, Switzerland, and Cambridge, U.K. in collaboration 
with the United Nations Environment Programme, and with financial support from the World 
Wide Fund For Nature. 

A CONTRIBUTION TO GEMS - THE GLOBAL ENVIRONMENT MONITORING SYSTEM. 



U Co) <tt 



Copyright 



1987 International Union for Conservation of Nature and Natural 
Resources/United Nations Environment Programme/World Wide Fund for 
Nature. 



This publication may be reproduced in whole or in part and in any form for 
educational or non-profit purposes, without special permission from the 
copyright holders provided acknowledgement of the source is made. 

No use of this publication may be made for resale or other commercial 
purpose, without the prior written permission of the copyright holders. 



Citation: 



IUCN/UNEP/WWF (1987). Madagascar, an environmental profile, 
by M.D. Jenkins. IUCN, Gland, Switzerland and Cambridge, U.K. 



Edited 



ISBN 



No 2-88032-607-9 



Printed by 



Unwin Brothers Ltd, The Gresham Press, Old Woking, Surrey. 



Cover Design James Butler 

Cover Photos LANDSAT image of north-west Madagascar showing Nosy Be and adjacent 

coast: NASA/UNEP. 

Baobab Adansonia za: M. Pidgeon. 

Rice paddies at Eminiminy, south-east Madagascar: S. O'Connor. 

Indri Indri indri: WWF/J.J.Petter 



Available from: 



IUCN Publications Services, 

219c Huntingdon Road, Cambridge CB3 ODL, U.K. 



The designations of geographical entities in this book, and the presentation of the material, do 
not imply the expression of any opinion whatsoever on the part of IUCN, UNEP and WWF 
concerning the legal status of any country, territory, or area, or of its authorities, or concerning 
the delimitation of its frontiers or boundaries. 



CONTENTS 

Page number 



IX 



Introduction 

Acknowledgements x 

A note on place names x 

Maps x j 

PART I. PHYSICAL GEOGRAPHY ] 

1.1. Geology 1 

1.2. Relief 2 

1.3. Soils 4 

1.4. Climate 5 

Temperature 5 

Rainfall 5 

Bioclimates 6 

Cyclones 7 

1.5. Hydrography 7 

River systems 7 

Lakes 8 

Hydrology 9 

PART II. HUMAN GEOGRAPHY 1 1 

II. 1. Population estimates and growth rates 11 

11. 2. Distribution of population 11 

11. 3. Ethnic divisions 1 1 

11. 4. Agriculture 14 

Land use and farm structure 15 

Crop production 16 

Livestock 20 

PART III. FLORA, VEGETATION AND FOREST COVER 23 

111.1. Flora 23 

Diversity 23 

Endemism 23 

Affinities of the flora 24 

Origins of the flora 25 

111. 2. Vegetation 27 

Eastern region 27 

Western region 29 

Secondary formations 30 

111. 3. Forest cover and destruction 32 

Eastern forests (inc. Sambirano domain) 32 

Western forests 33 

Southern forests 35 

Central highlands 35 

Montane vegetation 36 

111.4. Forest exploitation 36 

Timber 36 

Other forest products 37 

111. 5. Reafforestation 37 

111.6. Ethnobotany 38 

Introduction 38 

The potential value of the Madagascan flora 38 

Previous studies of ethnobotany in Madagascar 40 



-v- 



An environmental profile of Madagascar 

PART IV. MARINE AND COASTAL ECOSYTEMS 47 

IV. 1. Relief 47 

IV. 2. Mangroves 47 

IV. 3. Coral reefs 48 

Distribution of reefs 48 

Conservation of reefs 50 

Grand Recif proposed Marine National Park 51 

Nosy Be 54 

Offshore sand cays 55 

PART V. FAUNA 59 

V.l. Birds 60 

V.2. Mammals 63 

V.3. Amphibians and reptiles 66 

V.4. Fishes 72 

V.5. Lepidoptera: rhopalocera (butterflies) 77 

V.6. Terrestrial molluscs 80 

V.7. Freshwater molluscs 87 

V.8. Marine molluscs 90 

V.9. Nonmarine crustaceans 96 

V.10. Marine crustaceans 99 

V.l 1. Other invertebrates 106 

PART VI. PROTECTED AREAS 109 

Introduction 109 

Legislation 109 

Administration 109 

Total area under protection 110 

Information sheets 111 

Montagne d'Ambre National Park 112 

Isalo National Park 115 

Betampona Natural Reserve 116 

Zahamena Natural Reserve 118 

Tsaratanana Natural Reserve 121 

Andringitra Natural Reserve 123 

Lokobe Natural Reserve 126 

Ankarafantsika Natural Reserve 128 

Tsingy de Namoroka Natural Reserve 131 

Tsingy de Bemaraha Natural Reserve 132 

Lake Tsimanampetsotsa Natural Reserve 134 

Andohahela Natural Reserve 137 

Marojejy Natural Reserve 140 

Ambohitantely Special Reserve 144 

Beza Mahafaly Special Reserve 145 

Nosy Mangabe Special Reserve 148 

Perinet-Analamazoatra Special Reserve 149 

Analabe Private Reserve 152 

Berenty Private Reserve 156 

PART VII. OTHER IMPORTANT AREAS 157 

Rain forest areas 157 

Forests of Maroantsetra 157 

Masoala Peninsula 159 

Sihanaka Forest 160 

Ranomafana 160 

Non rain forest areas 162 

Ankarana Massif / Ambilobe Karst 162 

Ankaratra Massif 163 

Lake Ihotry 164 

Zombitse Forest 164 



Contents 

APPENDIX 1. ENVIRONMENTAL LEGISLATION 167 

A. Species legislation 167 

B. International agreements 168 

APPENDIX 2. FAUNAL LISTS 169 

Birds 170 

Mammals 178 

Reptiles 181 

Amphibians 188 

Fishes 193 

Butterflies (except Hesperidae) 197 

Nonmarine molluscs 204 

Nonmarine crustaceans 213 

APPENDIX 3. SPECIES ACCOUNTS 217 

A. Birds 218 

Tachybaptus pelzelnii 218 

Tachybaptus rufolavatus 220 

Ardea humbloti 221 

Anas bernieri 223 

Aythya innotata 225 

Haliaeetus vociferoides 226 

Eutriorchis astur 228 

Mesitornis variegata 230 

Mesitornis unicolor 231 

Monias benschi 233 

Sarothrura watersi 234 

Amaurornis olivieri 236 

Charadrius thoracicus 237 

Coua delalandei 238 

Tyto soumagnei 239 

Brachypteracias leptosomus 240 

Brachypteracias squamiger 242 

Atelornis crossleyi 243 

Uratelornis chimaera 245 

Neodrepanis hypoxantha 246 

Phyllastrephus apperti 248 

Phyllastrephus tenebrosus 249 

Phyllastrephus cinereiceps 250 

Xenopirostris damii 251 

Xenopirostris polleni 252 

Monticola bensoni 254 

Crossleyia xanthophrys 255 

Newtonia fanovanae 257 

B. Mammals (lemurs) 263 

Allocebus trichotis 263 

Cheirogaleus major 264 

Cheirogaleus medius 266 

Microcebus coquereli 268 

Microcebus murinus 269 

Microcebus rufus 271 
Phaner furcifer 

Avahi laniger 274 

/«rfr; /'«drj 276 

Propithecus diadema 278 

Propithecus verreauxi 279 

Daubentonia madagascariensis 282 

Hapalemur griseus 284 

Hapalemur simus 286 



Lemur catta 



288 



-Vll- 



An environmental profile of Madagascar 

Lemur coronatus 290 

Lemur fulvus 291 

Lemur macaco macaco 295 

Lemur macaco flavifrons 297 

Lemur mongoz 298 

Lemur rubriventer 301 

Lepilemur ruficaudatus 303 

Lepilemur dorsalis 304 

Lepilemur ed ward si 310 

Lepilemur leucopus 305 

Lepilemur mustelinus 307 

Lepilemur microdon 308 

Lepilemur septentrionalis 309 

Varecia variegata 3 1 1 

C. Reptiles 314 

Geochelone radiata 314 

Geochelone yniphora 3 1 7 

Pjuris planicauda 320 

Ppjcis arachnoides 322 

Erymnochelys madagascariensis 323 

Eretmochelys imbricata 325 

Chelonia mydas 326 

Lepidochelys olivacea 327 

Caretta caretta 327 

Crocodylus niloticus 328 

D. Lepidoptera 330 

Papilio grosesmilhi 330 

Papilio morondavana 331 

Papilio mangoura 332 

APPENDIX 4. PLANT SPECIES LISTS 335 

Succulents 336 

Palms 340 

APPENDIX 5. ETHNOBOTANY DATABASE 341 

Taxonomic table 342 

Medicinal usage table 357 

References 372 



-Vlll- 



INTRODUCTION 



Madagascar, with its wide range of natural ecosystems, unique and varied fauna and flora, 
largely rural human population and often severe environmental problems, is universally 
recognized as a high priority for conservation action. This report aims to provide information 
on which decisions affecting the environment of the country can be made, and to give 
indications of areas in which further research is necessary. It is essentially bibliographic in 
nature and represents a distillation of available sources of information, both published and 
unpublished. Bibliographies and reference lists are provided for each section. 

The principal subject areas covered are physical and human geography, vegetation, forest cover 
and loss, ethnobotany, marine and coastal ecosystems (in particular coral reefs), fauna, 
protected areas and sites of biological importance. With respect to fauna, emphasis has been 
laid on endemic and threatened taxa. All native vertebrate groups are discussed, but the sheer 
number and variety of invertebrates, particularly arthropods, has precluded their being treated 
in a similar fashion - it is evident that any attempt to cover all invertebrate groups in a single 
volume would result in extremely superficial treatment. Discussion has thus been confined to 
groups identified as of particular interest, notably crustaceans, molluscs and butterflies 
(Rhopalocera). 

Work on the volume began in 1983 and has continued intermittently until the present (early 
1987), with the most recent incorporation of new information being in January 1987. 

The report was compiled and edited by Martin Jenkins and authored by the following: 

Nigel Collar, Mark Collins, Tim Dee, Stephen Droop, Diana Evans, Brian Groombridge, Jerry 
Harrison, Martin Jenkins, Jane Thornback, Sue Wells, Lissie Wright. 

The sections on Ethnobotany (Part III.6 and Appendix 5) are taken from: Ethnobotany in 
Madagascar. Overview/ Action Plan/ Database. (1985). A report to WWF/IUCN prepared by 
Mark Plotkin, Voara Randrianasolo, Linda Sussman and Nina Marshall. 



-IX- 



ACKNOWLEDGEMENTS 



Many people have helped in many ways in the compilation of this report, by provision of 
information, by advice and by commenting on and reviewing parts of the text. In particular 
we would like to thank the following: R. Albignac, L. Allorge, J. Andriamampianina, C. Blanc, 
R. Blommers-Schlosser, Q. Bloxam, P. Bouchet, P. Brinck, D.S. Brown, A.C. van Bruggen, E.R. 
Brygoo, C. Carter, A. Crosnier, D. Curl, J. Dransfield, L. Durrell, B. Dussart, E. Fischer, D.G. 
Frey, J.T. Hardyman, H.H. Hobbs, L. Holthuis, A. Jolly, M. Keech, A. Kiener, P. Lake, O. 
Langrand, K. MacKinnon, R. Mittermeier, J and N. Moore, J. Moreau, M. Nicoll, S. 
O'Connor, R. Paulian, J. -J. Petter, M. Pichon, M. Pidgeon, J. Pollock, H.D. Rabesandratana, B. 
Rakotosamimanana, C. Raxworthy, A. Richard, Y. Rumpler, B. Salvat, P. Thompson, S. Tillier, 
B. Vaohita, B. Verdcourt, P. Viette, D. Vukidanovic, W. Weiss, J. Wilson, P. Wright. 

Errors and omissions remain the responsibility of the editor. 



A NOTE ON PLACE NAMES 



Seven of the principal port-towns of Madagascar, along with the capital and He Sainte-Marie, 
have recently changed their names; in almost all the available literature the old, French names 
are used. Every effort has been made in this report to use the current names, though some 
discrepancies may exist. 

New Old 

Antananarivo Tananarive 

Antseranana Diego-Suarez 

Fenoarivo Atsinanana Fenerive 

Mahajanga Majunga 

Nosy Borah He Sainte-Marie 

Taolanaro Fort Dauphin 

Toamasina Tamatave 

Toliara Tulear 

Vohimarina Vohemar 

There is also some variation in the spelling of other place-names (e.g. Marojejy/Marojezy, 
Ihotry/Iotry, Sihanaka/Sianaka, Analamazoatra/Analamazaotra), though there should generally 
be no possibility of confusion. Finally, while the Malagasy term for 'island 1 has generally been 
spelt in the currently accepted form 'Nosy' (e.g. Nosy Be, Nosy Mangabe), in some instances 
the alternative Nossi- or Nosi- has been used; again there should be no possibility of confusion. 



-x- 





45 


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Map 1 as A 




Nosy Be /Sjj|fo J \ 




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MOZAMBIQUE /P<T J 


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V — s^ ^-^^ /"Q / 

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-^7 /* Fianarantsoa / 

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MADAGASCAR 


25°S 


Iv. .<*<£/ TaolanaroV 


Principal towns, rivers & reefs 

150, km 

i i 









-XI- 



Map 2 



VEGETATION OF 

MADAGASCAR- 

after White (1983) 




VEGETATION KEY 



1b Evergreen 

Rainforest 
7 Dry deciduous 

Forest 
HbMosaicof rainforest 

and secondary grass 
22bMosaic °* deciduous 

forest and grassland 
5 Moist montane forest 

18 Secondary vegetion 

of highlands 
19c Undifferentiated 

montane forest 

41 Dry thorn scrub 

46 Mosaic of thorn scrub 
and secondary grass 

' ' Mangroves 



-XJ1- 



Map 3 



PROTECTED AREAS OF 

MADAGASCAR, . tdW 

Foret d Amt 

Forest cover c. 1950 



'Ambre 
Anolamerana 




n be 



gabe 



KEY 

ntegral Reserve 
O Special Reserve 
Forest cover 



Cap Sainte Marie 



100 200 300 400km. 
_i i i 1 



-Xlll- 



PART I. PHYSICAL BACKGROUND 

The island of Madagascar extends from 11°57'S to 25°35 , S and 43°14'E to 50°27'E in the 
Indian Ocean; it is separated from the African continent by the Mozambique Channel, only 
some 300 km wide at its narrowest point. With a north-south length of 1600 km and a 
maximum width of 580 km, Madagascar has a surface area of 587 000 sq km and is the fourth 
largest island in the world, after Greenland, New Guinea and Borneo. Relief is complex and 
variable though nowhere rises higher than 3000 m and climate, though also highly variable, is 
predominantly tropical, despite the southern part of the island extending below the Tropic of 
Capricorn. 



1.1. GEOLOGY 

Four main geological features of Madagascar have combined to produce the island's relief; 
these are: 

i. A Precambrian basement covering the eastern two-thirds of the island (excepting some 
small coastal areas which are sedimentary in origin). 

ii. A sedimentary region along the west coast, dating from Permian to recent. 

iii. Widespread volcanic intrusions, mainly Upper Cretaceous and secondarily 
late-Tertiary/Quaternary. 

iv. An extensive lateritic mantle, principally overlying the Precambrian basement. 

i. The Precambrian basement is much folded and entirely metamorphic; it outcrops over the 
eastern two-thirds of the island, an area of some 400 000 sq km. Knowledge of the mineralogy 
and petrography of the basement is generally reckoned good, though its stratigraphy and 
structure are still relatively poorly understood. The basement has been divided into three 
'systems', or stratigraphic subdivisions, originally based primarily on the distribution of useful 
minerals: 

a. The Androyan system, covering some 55 000 sq km in the extreme south, characterised 
by intense metamorphism, a prevalent sedimentary origin, a high frequency of magnesian and 
calcic rocks and the widespread presence of workable deposits of phlogopite mica and 
thorianite. The system apparently has a very rich assembly of highly metamorphic rocks and 
minerals. 

b. The Graphite system, which overlies the Androyan. This extends for some 
250 000 sq km, or most of the Precambrian basement and nearly half the land area of 
Madagascar. It is characterised by the widespread presence of graphite gneiss but is very 
complex and is regarded as the least well known of the systems, the category acting as 
something of a general repository. Metamorphism is average to strong, rarely intense, though 
almost the entire system has been migmatised which further hampers stratigraphic study. It 
outcrops from sea level to 2600 m. 

c. The Vohibory system. This covers a total area of around 55 000 sq km, split into 
relatively small areas scattered over the Graphite system. It is characterised by moderate to 
slight metamorphism and by the marked importance of metamorphosed basic volcanic 
formations. 

ii. The sedimentary region consists principally of slightly folded sedimentary formations along 
the west coast, west of the Precambrian basement. These form a continuous zone 30-200 km 
wide. The oldest strata are Permian and above these are Jurassic and Cretaceous sediments of 
very variable facies. Much of this belongs to the continental sedimentary system known as the 
Karroo system (essentially equivalent to the Karroo in southern Africa), starting in the Lower 
Permian and continuing to the Middle Jurassic, though with many marine incursions (often 



-1 



An environmental profile of Madagascar 

forming massive reef limestones and marls). The Upper Jurassic and Cretaceous formations are 
largely marine. 

Tertiary marine deposits outcrop in the Antseranana region in the north and then along the 
coast in a discontinuous strip from the Ampasindava peninsular to the Menarandra in the 
south. Superficial recent continental or marine deposits, sometimes very extensive, overlie 
much of this. 

The sedimentary region does not rise over 1300 m and is generally of gentler relief than the 
basement area. 

iii. Volcanic formations. The two main periods of volcanic activity are Upper Cretaceous and 
late-Tertiary/Quaternary. The most extensive volcanic formations are Upper Cretaceous, 
found both in the Precambrian basement and the sedimentary areas where they can be dated by 
their relationships to identifiable sedimentary layers; they derive from extensive basaltic flows 
(from fissure-type volcanic activity) and locally from sheets of rhyolitic ignimbrites. The later 
volcanic intrusions are of much lesser extent and are often difficult to date precisely as they 
occur largely in the Precambrain basement area; however they play a locally important role in 
relief and are discussed in I.2.iv below. 

iv. Weathering residues. Most of the Precambrian basement is mantled by a layer of lateritic 
clays or clayey laterites (produced by chemical weathering of plutonic, metamorphic or 
sometimes volcanic feldspars). Strictly speaking these are soils, but they are often so thick 
(10-15 m, sometimes up to 60-80 m) that they play an important geological and groundwater 
role. Laterisation has probably been going on since the Pliocene and is one of the principal 
causes of the accelerated erosion which is a major environmental problem in the country. 



1.2 RELIEF 

The island shows an extremely varied and often rugged topography, although it does not rise 
above 3000 m in elevation, (the highest point, Mt. Maromokotra in the Tsaratanana Massif, is 
2876 m). The main axis of the island extends in a north-north-east to south-south-west 
direction, with the main lines of relief along this, though there is marked asymmetry about the 
central axis. 

The eastern coastline, which is almost straight for a large part of its length (some 650 km), 
gives way to a narrow but continuous coastal plain. From this an escarpment (occasionally 
double) rises to a plateau of between 800 and 1500 m altitude which slopes down to the north 
and south and has many massifs of up to 2500 m altitude rising out of it. The westward slope 
is much gentler, though interrupted by cliffs, with the western regions consisting of plains and 
plateaux with extensive delta areas. 

A division can be made between the relief of the Precambrian basement and that of the 
sedimentary region to the west. 

i. The Precambrian basement has been considerably uplifted and faulted; this and the 
subsequent erosion explain the great variety of landscape forms. In general the gneiss outcrops 
have been profoundly affected by lateritic weathering and have formed a jumble of hills on 
which the characteristic 'lavakas' 1 form as a result of accelerated erosion. The granites and 



The term 'lavaka' is applied to the most prominent accelerated erosion forms which develop 
in the deep laterites of the Precambrian basin; these generally have the form of fan-shaped 
openings or cirques in hillsides. Individual lavaka may coalesce to form compound lavaka 
which may be several tens of metres deep and several hundred metres wide (for further 
discussion see Le Bourdiec, 1972). 



Physical background 

quartzites which are much more resistant to erosion usually form the high points of the 
landscape, often as rounded massifs such as Andringitra, which contains the second highest 
summit on Madagascar. Also important in this region are the 'tampoketsa' which are the 
levelled off remains of ancient erosion surfaces at high elevation, generally forming plateaux 
bordered by steep escarpments. The most important of these are in the north-west: at 
Fenoarivo, Ankazobe, Kamoro, Beveromay and Anaiamahitsy; these are assumed to be 
late-Cretaceous in origin (see below). 

ii. The sedimentary region in the west contains the two main sedimentary basins on the island 
- Mahajanga and Morondava, one south, the other north of Cap Saint Andre. The region 
consists mainly of alternating hard and soft beds dipping gently towards the sea (normally at an 
angle of some 3 to 5°, occasionally decreasing to 2° and rarely exceeding 10°). Erosion has led 
to the formation of a landscape of cuestas. In the sandstone cuestas such as the Isalo in the 
south-west, the reverse (steeper) slopes tend to be dissected by a labyrinth of deep canyons or 
to be cut into ruiniform relief. The limestone cuestas tend to be subjected to karst processes, 
often leading to a highly dissected, block-like topography (as in parts of the Bemaraha and 
Ankarana karsts) and the presence of dolines and extensive cavern systems. There are 
estimated to be some 33 000 sq km of karst, the most important of these being: Bemaraha; 
Kelifely plateau; Ankara plateau; Sitampiky; Ankarana; Mahafaly; Mahajanga and the Narinda 
peninsula. 

Two other factors contribute to the landscape across both regions: 

Hi. Erosion surfaces. The earliest of these, attributed to the late-Cretaceous, affects the 
Precambrian basement and has been alluded to above. Two others of importance have been 
identified, corresponding to the mid- and late-Tertiary. The former, below the level of the 
late-Cretaceous surface, is believed responsible for many of the medium-sized hills in the 
landscape in the central plateau, particularly around Antananarivo; it has also levelled off parts 
of the sedimentary cover in the west: the summit plateaux between 900 and 1000 m in the 
sandstone Isalo massif are thought to result from it. The late-Tertiary erosion cycle has 
resulted in vast pediplains in both the basin and sedimentary areas, for example north of the 
Androy and Mahafaly massifs in the far south, the Zomandao plain and the reverse slope of 
the Isalo massif. It also appears at the base of the principal basins of the central highlands, 
where it has been dissected into a system of small hills of 50 to 100 m relative height. 

iv. Volcanic forms. These constitute many of the major massifs on the island and occur 
principally in the following areas: 

a. In the north the massifs of Ankaizina and Tsaratanana. The latter is the highest massif 
on Madagascar; both areas have necks of phonolite, with relief determined by differential 
erosion. The Ankaizina also has trachytes and two series of recent basalt eruptions, the first 
heavily eroded, the second well-preserved with many small volcanic cones, some with crater 
lakes. 

b. The Itasy and Ankaratra massifs in the central highlands around Antananarivo; these are 
the most extensive volcanic regions. The Itasy is a 750 sq km assemblage of cones and domes 
with well preserved forms, produced by volcanic emissions in a gneissic depression; most of the 
domes are below 1600 m, though the gneiss is often higher than this. The Ankaratra is much 
larger (some 4000 sq km) and can be divided into three main regions: the first, in the 
north-east, consists of a strong line (running NNE-SSW) of volcanos dating from the end of the 
Pliocene or early quaternary with compact lava (ankaratrites); several summits exceed 2400 m 
(the Tsiafajavona is the third highest massif on island, reaching 2643 m). The second is in the 
western and central southern region of the massif, consisting of old rhyolitic and trachyitic 
Pliocene domes on which are superimposed huge, more recent, basalt flows which have resulted 
in barrage lakes and waterfalls on the watercourses. The third, situated in the south and 
south-west of Betafo and Antsirabe, represents the most recent volcanic activity, with 
numerous broken cones and crater lakes. 



An environmental profile of Madagascar 

c. The Androy in the far south consists of a stack of superimposed flows, alternating basalt 
and rhyolite, dating from the end of the Cretaceous; relief results from differential erosion, 
with the basalt eroding easily, while the rhyolites have formed a large cuesta which encircles 
the massif (reaching 600 m). In the centre of the massif is a table of subhorizontal rhyolites of 
700 m altitude. 



1.3. SOILS 

As with relief, there are marked differences between the soils of the Precambrian basement 
and the western sedimentary region. 

i. The Precambrian Basement. Over the great part of the Precambrian basement, the soils are 
composed of lateritic clays; these are found in both forested and savanna regions and have 
virtually no surface humus. Hydroxides of aluminium and iron in them are mixed with a high 
proportion of clay and there is little surface concretion. Studies (cited in Guilcher and 
Battistini, 1967) have shown that although these soils are poor, they are not absolutely infertile, 
though leaching has led to impoverishment of alkaline and alkaline earth elements, and of 
silicates. Lateritic clays are the soils currently under formation in most areas. There are also 
far less widespread lateritic hardpans which are Pliocene or earlier in origin; these are 
considerably more prone to laterisation than the recent clays. 

Above 2000 m, the clays are replaced by shallow grey arenaceous soils, or in the small basins 
of the quartzite crests by quartz sands. In the south and south-west of the Precambrian basin 
are found soils particular to the region: one is a soil of the calcareous crust, especially on 
gneiss, where cipolines and amphibolites provide calcareous elements for the soil (these can be 
cultivated if adequately irrigated); the others are variable red non-lateritic soils, sometimes 
skeletal and leached, sometimes deep and rich. There is a very sharp dividing line between 
these and the characteristic lateritic clays west of the principal Anosyenne chain in the 
south-west of the island. 

The lateritic clays, which are essentially climatic soils, are found almost entirely over the 
crystalline and metamorphic rocks of the Precambrian basement; far fewer lateritic soils are 
found on most of the recent volcanic rocks. In the Itasy and Ankaratra massifs are found 
mostly black soils very rich in humus; these are analagous to the chernozems found in valley 
bottoms in the Androy in the western sedimentary region where they are derived from basalts 
(the rhyolites in the higher reaches do not produce them). These are relatively rich soils, 
whether they are forested or denuded. 

Alluvions are also found in the Precambrian basement region; their fertility is a function of 
their consituent elements and their age. They are mainly developed on the high plains, and 
those of the Ankaizina, for example, are generally rich. However the older alluvions, which 
are usually found at higher levels in drainage basins, have normally undergone laterisation and 
are consequently impoverished; this is particularly notable in the region of the Alaotra basin. 
The recent alluvions of the lower regions are often degraded by massive influxes of sandy 
elements from leaching of the lateritic clays originating in lavakas in the surrounding areas 
(e.g. in the Antanetibe plain in the upper Betsiboka). 

All in all the soils of the Precambrian basin are of average quality for tropical soils; they are 
often deficient in phosphoric acids but their nitrogen content is often good. However, all these 
soils except the lateritic and calcareous hard-pans are highly erodable, especially the lateritic 
clays. 

ii. The sedimentary region. There is a wide variety of soils in the western sedimentary region, 
though the two most notable features are the virtual absence of true lateritic clays and the very 
wide surface cover of a 'sand-clay carapace' (carapace does not imply hardness, but simply an 
overlying of the bedrocks) - this latter is a collective term which covers a considerable variety 
of conditions. 



-4- 



Physical background 

Although there are no true lateritic clays, there are three types of red soils: decalcified clays on 
calcareous rocks (terra rossa), found widely on the limestone karsts; red soils on cretaceous 
basalts; reddish soils of the semi-arid regions, notably silicaceous sands coloured by iron 
hydrates, found in the Androy region (these are not necessarily infertile). 

The marls and clays of the west never laterise. They form grey soils which have essentially the 
same composition as the bedrock, though with a higher organic content. In the recent volcanic 
massifs (Nosy Be, Mt d'Ambre) are found the same humus-rich dark soils as in the recent 
volcanic regions of the Precambrian basement. 

The sand-clay carapace, often very thick, can be a soil or a superficial geological formation 
according to circumstances; it covers large areas of a variety of rocks, especially sandstones or 
their surrounds. It is principally found in littoral or sublittoral zones, especially in the 
southern part of the island, though is also found in the interior, notably south of the Mangoky 
river. 

There are also soils of the recent alluvial plains, notably in the large western deltas. 

Overall, deforestation has led to somewhat less active erosion than in the Precambrian 
basement. This is doubtless because of the much higher frequency of flat or nearly flat areas 
(especially in the immense sub-littoral plains from Cap Saint Andre to the extreme south, but 
also elsewhere) and the permeability of the sand-clay carapace and of the karst plateaux (when 
the latter are not covered by terra rossa). However, although lavakas tend not to form, several 
erosive features which may have analagous effects are found in areas of deforestation - e.g. 
gully erosion on the reverse slope of the Isalo cuesta, between Ranohira and Sakaraha, and 
hemicyclic erosion in red soils on basalt, such as on the Radama peninsula - the latter implies 
that deforestation may have as drastic effects on red soil basalts as on the lateritic clays. 



1.4. CLIMATE 

Temperature 

In the lowlands the mean annual temperature is fairly uniform, though decreases from north to 
south, ranging from ca 27°C (Antseranana) to 23°C (Taolanaro) with mean annual range 
increasing from around 3°C in the north to 7.5°C in the dry south-west region. In the west 
this is often masked by local conditions and moderated by the Mozambique channel, a warm 
sea with very little circulation, and the Fohn effects of the trade winds descending from the 
central plateau. 

Altitude has a significant effect on temperatures, with an average lapse rate of around 0.6°C 
for every 100 m change in level. Generally lapse rate is below average on the eastern slopes, 
where the high humidity damps out the temperature variations, and above in the west. 

On the central plateau, mean annual temperatures usually lie between 16° and 19°C. Frost 
sometimes occurs above 1500 m though snow is virtually (but not completely) unknown. The 
mean annual range is between 5° and 6°C in the north and 7°C in the south; diurnal range is 
greater, being for example between 6° and 16°C in Antananarivo (1381 m., latitude 18°56'S). 

Rainfall 

Rainfall in Madagascar is governed by a double gradient: the annual amount decreases from 
east to west and from north to south, while seasonality increases in the same directions. Thus 
travelling westward and southward the dry season becomes longer and more marked - in the 
extreme south-west the climate is sahelian or semi-desert while the eastern coastal area is a 
subequatorial region of high rainfall and humidity throughout the year. Where seasonality is 
marked the dry and cool seasons are coincident (roughly June - October), making the whole 



An environmental profile of Madagascar 

island characteristically tropical in climate despite the south being below the Tropic of 
Capricorn. 

The major factor affecting rainfall is the interaction of the south-east trade wind (the 
predominant wind at all seasons), produced by the Indian Ocean anticyclone, with the direction 
of the principal lines of relief on the island (running NNE-SSW). North and north-west 
'monsoon' air currents which are a continuation of the north-east trade wind exert an 
important secondary effect. 

Orographic ascent, along the eastern coast and escarpment, of unstable moisture-laden air 
carried by the south-east trade wind leads to extensive cloud formations and heavy rainfall in 
this region. In summer the trade wind is somewhat attenuated by the withdrawal of the Indian 
Ocean anticyclone to the south and east, the effect being much more marked in the north than 
in the south. 

Above the moist convective trade wind layer there is a much drier stable air mass whose lower 
level is indicated by a subsidence inversion, also moving westwards. This inversion is most 
marked and at its lowest altitude in winter, from July to September, and at this time limits 
cumulus cloud formation and rainfall in areas away from the eastern coast. This combines with 
warming and drying Fohn effects of the trade wind descending the western slope of the 
plateau, to lead to generally dry and clear weather in western and southern parts in winter 
when the south-east trade wind dominates weather conditions. 

In summer a zone of intertropical low pressure affects the island and brings with it north or 
north-westerly 'monsoon' air currents, which are a continuation of the north-east trade wind. 
These are also heavily moisture-laden and bring a large amount of rain, though their effect 
decreases markedly eastwards and southwards. 

In winter the intensification of the Indian Ocean anticyclone and consequent south-east trade 
winds tends to drive the intertropical low pressure zone and monsoon air away to the north, 
considerably reducing the rainfall from this source. 



Bioclimates 

Rainfall and temperature variations combine to produce a number of 'bioclimatic regions' on 
the island. The boundaries between these are to some extent arbitrarily designated, though it is 
worth noting that in parts of the island (such as the south-east) the climate can change 
dramatically in character over the distance of a few kilometres. 

a. On the east coast, the climate is subequatorial with rainfall exceeding 1500 mm and 
sometimes 3000 mm, with no ecologically dry month and high temperatures throughout the 
year. 

b. On the eastern slopes of the plateau, the rainfall still exceeds 1500 mm with a dry season 
of 1 to 4 months and a mean temperature in the coldest months of between 10° and 15°C. 

c. On the western slopes of the plateau the dry season is longer, up to 5 or 6 months, 
rainfall is likely to be less than 1500 mm, though humidity in the dry season is still high 
through the dominating effects of the trade wind at this time of year. 

d. On the western plains, the dry season lasts 7 or 8 months in southern regions, somewhat 
less further north. Rainfall is around 1500 mm in the north, 500 to 1500 mm in the south. 

e. In the extreme south, rainfall is sparse and highly irregular in occurrence, being stormy 
and very localised. Some areas may not have any precipitation for 12 to 18 months. There is 
an absence of detailed climatic data for this region, making characterisation particularly 
difficult. 



Physical background 

These divisions correspond well with Humbert's phytogeographical divisions of the island based 
largely on the concept of a natural climatic plant climax community. 



Cyclones 

A factor of some considerable environmental importance in Madagascar is the prevalence of 
cyclones. Donque noted in 1972 that since 1848, 155 cyclones had hit the island, representing 
just under one quarter of all cyclones in that time in the south-western Indian Ocean. They 
occur in summer only, with the great majority between mid-January and mid-March; most hit 
the island along the north-eastern coast, travelling south-west and curving south-east across the 
island, though others hit the island from the Mozambique channel. They can cause great 
devastation, with winds of up to 300 km/hr and rainfall of 600 to 700 mm in four or five days, 
often resulting in large-scale flooding and massive destruction of crops and forests. 



1.5. HYDROGRAPHY 

River systems 

Following Aldegheri (1972), Madagascar is divided into five hydrographic regions of very 
unequal size: 

a. The slopes of the Montagne d'Ambre in the extreme north; 

b. The slopes of the Tsaratanana Massif; 

c. The eastern slopes which run into the Indian Ocean; 

d. The western and north-western slopes whose waters run into the Mozambique channel; 

e. The southern slopes. 

a. The Montagne d'Ambre region covers only some 11 200 sq km or barely 1.8% of the land 
area of the island, in the extreme north. The volcanic massif is drained by narrow torrents 
with few tributaries running in beds littered with blocks of basalt. The major rivers are the 
Irodo, the Saharenana and Besokatra; the waters of the last are used to the supply the town of 
Antseranana. 

b. The drainage area of the Tsaratanana Massif covers around 20 000 sq km; rivers are 
characterized by having very steep gradients (30 or 40 m/km) in the upper reaches, levelling 
off to only a few m/km on the coastal plains, both on the western and eastern slopes. There 
are four major rivers: the Mahavary (160 km long), the Sambirano (124 km) and the 
Maevarano (203 km) all flow into the Mozambique Channel, while the Bemarivo (140 km) 
flows east into the Indian Ocean. 

c. The eastern slopes cover around a quarter of the land area, or ca 150 000 sq km, 
extending in a strip some 1200 km long and averaging 100 km wide (ranging from 50 km along 
Beampingaratra north of Taolanaro to 190 km at the latitude of the Ankaratra Massif). 

Watercourses are comparatively short, with steep profiles. Because of the prevalence of 
secondary hill chains running parallel to the coast, linked by faults perpendicular to the coast, 
the watercourses are often many times longer than the direct distance from source to coast. 
The rivers meander in the narrow coastal plain and feed a chain of lagoons separated from the 
sea by sand dunes. These lagoons have been artificially joined along the central part of the 
coast to form the 400 km Canal des Pangalanes. 

There are five main rivers in this region: the Mananara, Mangoro, Rianila, Maningory and 
Mananjary. The Mananara is the longest of these at 418 km (though the source is only 50 km 
from the coast) while the Mangoro is the largest in terms of size of drainage basin 
(17 175 sq km) and volume of water carried. The Alaotra basin (discussed in more detail 
below) feeds into the Maningory. 



-7- 



An environmental profile of Madagascar 

d. The western slopes cover almost 365 000 sq km or over 60% of the land area; 
watercourses here are divided by Aldegheri into two groups: large rivers which flood widely 
over the Hauts Plateaux, having roughly triangular drainage basins with the apex towards the 
coast; and coastal streams located between these whose sources are on the western edge of the 
Hauts Plateaux. 

In the first group seven major basins are identified including the five largest basins in 
Madagascar. From north to south they are: the Sofia with a drainage basin of 27 315 sq km 
and major tributaries the Anjobony and the Bemarivo; the Betsiboka-Mahajamba system which 
covers 63 450 sq km, making it the largest basin in Madagascar; the Mahavavy, covering an 
area of 16 475 sq km; the Manambolo at 13 970 sq km; the Tsiribihina at 49 800 sq km; the 
Mangoky, the longest river on the island (821 km) with a basin of 55 750 sq km; and the 
Onilahy at 32 000 sq km. 

The numerous small coastal rivers all have basins of less than 8000 sq km. 

e. The southern slopes are divided into three parts: 

- The Mandrare basin in the east, which covers some 12 570 sq km. The Mandrare River rises 
in the Beampingaratra Massif, running for some 270 km; it is the only Madagascan river with a 
longitudinal profile very close to the equilibrium profile. 

- Three rivers, the Manamboro, Menarandra and Linta make up most of the Androy region in 
the extreme south, although here there are also several closed basins with no outlet to the sea, 
the largest of these being the Ampamabora bowl north of Amborombe. 

- The Mahafaly plateau in the west which has virtually no surface water or rivers. 



Lakes 

Reiner (1963) has provided an inventory of waterbodies (lakes, lagoons and large ponds) in 
Madagascar of over 20 ha in extent, listing over 530. The majority of these are small (less than 
100 ha), and only 18 exceed 1000 ha in area. 

Of the five largest lakes, two are in the Hauts Plateaux region (Alaotra and Itasy) and three in 
the west (Kinkony, Tsimanampetsotsa and Ihotry). 

a. Lake Alaotra is the largest lake on Madagascar; its minimum extent (i.e. during the dry 
season) is some 22 000 ha, though during the height of the flood an additional 35 000 ha of 
marsh to the south and west are entirely under water. The lake is very shallow, having a 
maximum depth of around 2 m during the dry season and 4 m at highest water. The lake is at 
750 m altitude and is in an area which has suffered considerable deforestation and subsequent 
erosion; soil run-off results in the lake waters being highly turbid and brown in colour. 

b. Lake Kinkony (17°09'-18°04'S, 48°15'-48°40'E), the second largest lake, covers an area 
of ca 10 000 ha at low water and 14 900 ha at high water; maximum recorded depth at high 
water is 4 m. The lake lies in the basin of the lower Mahavavy and is formed from a natural 
alluvial impoundment. 

c. Lake Ihotry (2P50'S, 43°30'E) is a closed lake in the region of the lower Mangoky; it 
shows great variations in surface area, ranging from a minimum of 865 ha to around 9 400 ha. 
Salinity varies accordingly, being higher than that of sea water at lowest water but over ten 
times less than this at high water. Maximum depth is 3.8 m. 

d. Lake Itasy (19°07'S, 46°45'E) is a 3500 ha lake situated in the volcanic Itasy Massif in 
the centre of the island; it is not strictly a crater lake, but rather a lake blocked by a lava 
outflow. It lies at a mean altitude of 1221 m and has a maximum depth of 6.5 m. 

e. Lake Tsimanampetsotsa, situated on the edge of the Mahafaly Plateau, is a shallow, saline 
lake of ca 20 by 3 km (its surface area ranges from ca 1600 to 2900 ha), saturated with calcium 



Physical background 

and magnesium sulphates; in areas on the east shore freshwater rises to the surface. The lake 
forms part of the Reserve Naturelle Integrate de Tsiminampetsotsa (R.N.I. No 10) (see Part VI). 

Hydrology 

All rivers show high discharge, often in the form of sudden violent spates during the rainy 
season (November to March-April) and particularly after cyclones (mostly mid-January to 
mid-March). Response to rainfall is usually almost instantaneous and in many areas there are 
daily flood peaks during the rainy season, usually at night. 

In the north and east rainfall is high throughout the yea. - and low water dishcarge is 
correspondingly high; water flow is thus generally abundant throughout the year. Flood waters 
are violent, especially on the east coast, because of the rugged relief and the direct exposure to 
cyclones from the Indian Ocean. 

In the west a distinction can be drawn between large and small watercourses. There is very 
little rain in the dry season and at this time of year the volume of water in small watercourses 
usually decreases downstream, being gradually lost by seepage to the water table; most of these 
streams are thus dry in their lower reaches from April/May to November. Spate waters 
generally appear very suddenly and are of short duration (often only a few hours). The larger 
rivers, however, have catchment areas which extend over the Hauts Plateaux region; here there 
is usually some rain during the dry season which, combined with the relatively high retention 
of the predominant lateritic soils, ensures that low waters are generally well sustained, though 
still considerably lower than on the east coast. This effect carries over into the western region 
and these rivers thus continue flowing throughout the year. Dry season discharge rates are still 
much lower than those in the wet season, with a relative decrease from north to south as the 
extent and severity of the dry season increases - thus the Mangoky has specific low water 
discharge values about 10 times less than those of the Betsiboka some 600 km further north. 

In the south, with a very marked dry season and erratic rainfall in the wet season, rivers can 
show very rapid spate and extreme variations in water level - Aldegheri reports a case on the 
Menarandra of a rise of almost 3 m in water level in under 20 minutes. During the dry season 
flows decrease to such an extent that few rivers carry water as far as the sea throughout the 
year. However there is usually an underground supply in the riverbed sand which is used by 
local people. 



PRINCIPAL REFERENCES 

Aldegeheri, M (1972). Rivers and streams on Madagascar. In: Battistini, R. and 

Richard-Vindard, G. (Eds), Biogeography and ecology in Madagascar. Monographiae 

biologicae 21. Junk, the Hague. 
Battistini, R. (1972). Madagascar relief and main types of landscape. In: Battistini, R. 

and Richard-Vindard, G. (Eds), Biogeography and ecology in Madagascar. Monographiae 

biologicae 21. Junk, the Hague. 
Brenon, P (1972). The geology of Madagascar. In: Battistini, R. and Richard-Vindard, G. 

(Eds), Biogeography and ecology in Madagascar. Monographiae biologicae 21. Junk, the 

Hague. 
Donque, G. (1972). The climatology of Madagascar. In: Battistini, R. and 

Richard-Vindard, G. (Eds), Biogeography and ecology in Madagascar. Monographiae 

biologicae 21. Junk, the Hague. 
Guilcher, A. and Battistini, R. (1967). Madagascar - geographic regionale. 'Les cours de 

Sorbonne', Centre de documentation universitaire, Paris. 
Humbert, H. (1954). Les territoires phytogeographiques de Madagascar. Leur cartographie. 

Les divisions ecologiques du monde. CNRS 195-204. 
Humbert, H. and Cours-Darne, G. (1965). Carte intemationale du tapis vegetal, 

Madagascar. French Institute of Pondicherry. 



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An environmental profile of Madagascar 

Jolly, A., Oberle, P., and Albignac, E.R. (Eds) (1984). Key Environments - Madagascar. 

Pergamon Press, Oxford. 
Keiner, A. (1963). Poissons, peche et pisciculture a Madagascar. Centre Technique 

Forestier Tropical, Nogent-sur-Marne. 
Le Bourdiec, P. (1972). Accelerated erosion and soil degradation. In: Battistini, R. and 

Richard-Vindard, G. (Eds), Biogeography and ecology in Madagascar. Monographiae 

biologicae 21. Junk, the Hague. 
Moreau, J. (in press). Region 9. Madagascar. In: Burgis, M.J. and Symoens, J.J. (Eds), 

African Wetlands and Shallow water bodies. Vol 2. Directory, ORSTOM, Paris. 
Rossi, G. (1976). Karst et dissolution des calcaires en milieu tropical. Z.Geomorph. N.F. 

Suppl.-Bd. 26: 124-152. 



-10- 



PART II. HUMAN GEOGRAPHY 

II. 1. POPULATION ESTIMATES AND GROWTH RATES 

The most recent population census, in 1974/75, gave a figure of 7 603 790 (Thompson, 1982). 
UN mid-year population estimate for 1985 is 10 012 000. 

Growth rates are extremely high: the UN estimate for average growth rate over 1980-85 is 
2.8% per annum, giving a population doubling time of 25 years. In 1974/75 over half the 
population was under 20 years old (Thompson, 1982). 

In 1972 registered birth rate was 37.4/1000 and death rate 11.1/1000, though birth registration 
was estimated to be only 70% complete, death registration 50%, this giving real figures of 
53.4/1000 and 22.2/1000 respectively. 

II.2. DISTRIBUTION OF POPULATION 

Regional breakdown of population is given in Table 1. The population is very largely rural; 
Battistini and Verin noted in 1972 that 86% of people lived in villages of fewer than 2000, only 
14% of the population living in agglomerations of 2000 or over, and 8% in towns of over 
20 000. 

The population is very unevenly distributed over the island with population growth rates also 
varying regionally. Average density in 1981 was around 15 per sq km. Regions of higher 
population density are mainly in the central highlands and eastern coastal plain. Battistini and 
Verin (1972) noted that high local densities of over 50 per sq km (with patches of over 
150 per sq km) (presumably outside towns), were coincident with regions of intensive rice 
cultivation, mainly in the central highlands: around Antananarivo (Betsimitatatra region), 
Antsirabe (Vakinankaratra) and in the Betsileo country around Ambositra, Ambohimahasoa and 
Fianarantsoa, but also on the eastern coastal plain, notably in the Farafangana, Manakara, 
Mananjary and Fenoarivo Atsinanana regions. 

In large areas of the west and south, population densities are very low, from 2 to 5 per sq km., 
with the inhabitants largely pastoralists. Areas of shifting cultivation, such as much of the 
eastern escarpment slopes, and some rice growing areas (e.g. the Tsimihety highlands) have 
intermediate densities (5 to 15 per sq km). 

These variations and the widely differing land-use patterns in different parts of the island lead 
to marked regional differences in environmental impacts and problems. 



II.3. ETHNIC DIVISIONS 

The origins of the Madagscan people are complex and incompletely understood, though appear 
to be based on successive waves of migration from both Indonesia and Africa (most 
importantly the former), with Arab influences from the twelfth century onwards and contact 
with Europeans dating from the sixteenth century. There is no evidence for human occupation 
earlier than 2500 years ago. There is now essentially a single, though diverse, culture and a 
predominantly Indonesian language of which several mutually understandable dialects exist. At 
present, around 20 tribes are generally recognized, though these are based more on old 
kingdoms than on genuine ethnic groupings. Continuous migration and increased 
communication in the present century, along with a marked cultural unity, has tended to break 
down many geographic and ethnic barriers. However strong fidelity to traditional homelands 
persists, and the custom of endogamy remains widespread, militating against intertribal 
marriage. Historical tensions still manifest themselves, in particular between the peoples of the 
Hauts Plateaux, especially the traditionally elite Merina, and the coastal tribes, collectively 
known as 'coders'. 



•11- 



An environmental profile of Madagascar 



TABLE I. REGIONAL BREAKDOWN OF POPULATION (1978) 



Province 


Area Sq km 


Population 


Mean density 


Chief Town 


Population 


Antseranana 


42 725 


620 228 


14.5 


Antseranana 


48 000 


Mahajanga 


152 165 


857 610 


5.6 


Mahajanga 


57 500 


Toamasina 


72 212 


1 254 639 


17.4 


Toamasina 


59 100 


Antananarivo 


57 775 


2 322 109 


40.2 (33.3) # 


Antananarivo 


400 000 


Fianarantsoa 


100 326 


1 908 465 


19.0 


Fianarantsoa 


55 500 


Toliara 


162 283 


1 084 083 


6.7 


Toliara 


34 500 



Figure in parentheses excludes Antananarivo. 

Source: Bulletin mensuel de Madagascar (from 1971) continuation of the trimestrial Bulletin de 
statsitique generale de Madagascar (1949-71), Service de Statistique Generale, Antananarivo. 



TABLE 2. ETHNIC GROUPS IN MADAGASCAR (1972) 

Ethnic group Number 



Merina 
Betsimisaraka 


1 934 765 
1 106 991 


Betsileo 


892 352 


Tsimihety 


533 289 


Sakalava 


434 315 


Antandroy 


396 820 


Antaisaka 


377 110 


Tanala 


283 908 


Antaimoro 


255 161 


Bara 


250 261 


Sihanaka 


182 948 


Antanosy 


172 797 


Mahafaly 


120 620 


Antaifasy 


88 899 


Makoa 


80 069 


Bezanozano 


56 588 


Antakarana 


44 852 


Antambahoaka 


29 481 


Other Madagascans 


80 245 



Source: Area Handbook for the Malagasy Republic (1973). 



-12- 



Human geography 

Merina Inhabit the central highlands around Antananarivo, the city being 95% Merina; the 
region itself is called Imerina. The tribe is divided into three castes: Andriana (Nobles), Hova 
(Free-men), Andevo (descendants of former slaves). There is much rice cultivation by 
irrigation. The Merina formed the last and most powerful of the Madagascan kingdoms, which 
by the nineteenth century controlled most of the island; they were the first tribe to develop any 
skill in architecture and metallurgy and were the first to use a metal bladed 'angady', the 
long-handled Malagasy spade which is still the principal agricultural tool. 

Betsimisaraka The second largest tribe, living on the east coast in the Toamasina - Antalaha 
region. Some practice slash-and-burn cultivation on hillsides, growing mountain rice and 
maize, others practice irrigated high-density rice cultivation in scattered patches, often at the 
mouths of the principal rivers; there is also large scale production of cash crops, in particular 
coffee. The inhabitants of Nosy Borah (He Ste Marie) are sometimes considered Betsimisaraka. 

Betsileo Centred in the south of the Hauts Plateaux around Fianarantsoa but about 150 000 
live in the Betsiboka in the north-west (Mahajanga province). Much rice growing, often on 
hillside terraces, and some scanty pastureland. 

Tsimihety Descendants of the Sihanaka, inhabiting the north-central area, spreading west. 
Generally rice-growers, though they live in a region of low population density, with extensive 
thinly populated grazing lands between rice growing areas. Cattle are of very great Locial 
importance. 

Sakalava Occur in the west between Toliara and Mahajanga. The Sakalava were the first 
important Madagascan kingdom, founded at the end of the sixteenth century, but were largely 
conquered by the Merina in the nineteenth century. Essentially cattle raisers, with large herds 
grazing over enormous areas. Dry crops occupy a small amount of land around the villages. 
Near some of the rivers through the area there are considerable rice-growing areas, though 
these are apparently mainly the work of the Betsileo and Antaisaka immigrants south of Cap 
Saint-Andre, and the Betsileo and Merina on the lower Betsiboka. 60 000 inhabitants of the 
Mangoky delta, sometimes called 'Masikoro' are generally included in the Sakalava; the Veza 
fishermen (qv) are also sometimes considered Sakalava. 

Antandroy Nomadic, living in the arid south around Ambovombe. Dark skinned, different 
from other tribes. Primarily pastoral. Very little rice is grown, mostly millet, maize, cassava 
and beans. 

Antaisaka An offshoot of the Sakalava tribe, though now largely of mixed descent (e.g. with 
Bara and Tanala elements), centred south of Farafangana on the south-east coast. The poverty 
of the region and the high population growth rate has led to much migration, both seasonal and 
permanent. By 1970 nearly 40% of Antaisaka lived outside their homeland. They principally 
cultivate coffee, bananas and rice. 

Antaimoro Live in the south-east around Vohipeno and Manakara. They are of Islamic 
extraction and, unusually, have written records dating back to 1335, when they arrived on the 
island, they being one of the most recent peoples to arrive. Principally rice-growers. 

Bara Nomadic cattle-raisers, originating in the south-west near Toliara, they now live in the 
south-central area around Ihosy and Betroka. 

Tanala Forest people living inland from Manakara, occupying about half of the eastern 
mountain escarpment. They practice slash-and-burn but are also skilled hunters, gatherers and 
woodsmen. Increasingly, improved cultivation methods have been adopted, especially in the 
growing of coffee and the irrigation of rice. 

Sihanaka Inhabit the central plateau region north-east of the Merina, in the region around 
Lake Alaotra. They have much in common with the Merina and live principally by fishing, 
growing rice and raising poultry; they inhabit one of the best regions for agricultural 
development, with large scale rice growing projects taking place around Lake Alaotra. 



•13- 



An environmental profile of Madagascar 

Antanosy Live around Taolanaro in the south-east and are principally rice-cultivators. 

Mahafaly Inhabit the extreme south-west. Here, along the coastal strip, there are densely 
populated rural regions where almost all the land, especially that on the Karimolian dune sands, 
is occupied by contiguous plots of land ('vala') enclosed by aloe hedges; in these dry land crops 
such as manioc, sorghum, sweet potato and beans are grown. No rice is cultivated. Cattle 
raising is of secondary importance and is carried out inland on the crystalline pediplain in the 
north of the Mahafaly region. 

Antaifasy Live in the south-east around Farafangana; they cultivate rice and also practice 
fishing in lakes and rivers. 

Makoa Originally spread along the north-west coast, many have now moved south to the area 
of the Onilahy River. They are believed to be descended from African slaves and are the only 
true negroid peoples on the island. 

Antankarana An offshoot of the Sakalava dynasty, now a heterogenous group of mixed 
Sakalava, Betsimisaraka and Arab ancestry living in the north-west around Antseranana; 
mainly cattle raisers, they also grow dryland crops such as maize, rice and cassava. 

Bazanozano One of the first tribes to arrive, these live between the Betsimisaraka lowlands 
and the Merina highlands. They are predominantly herders and woodsmen, though they also 
grow some rice. 

Antambahoaka The smallest tribe, they are rice cultivators who live around Mananjary on the 
south-east coast. These are of the same Islamic descent as the Antaimora. 

Also recognized are: 

Vezo A clan of the Sakalava living in the west from Morondava to Faux Cap; they live 
by fishing and do not cultivate rice. 

Zafimaniry Live in ca 100 villages between the Betsileo and Tanala, are descended 
from high plateau people who migrated to the region early in the nineteenth century. 

Mikea These are hunter-gatherers whose existence as a separate ethnic entity has been 
questioned. They are found in the south-west, in the Mikea forest between Manombo 
and Morombe north of Toliary where they live in very tight groups of up to 15, having 
no contact with other tribes. Battistini and Verin aver that these people are 
Sakalava-Masikoro, also stating that on the Mahafaly plateau south of the Onilahy river 
there are Mahafaly with a 'Mikea' lifestyle. 

'St Marians' Live on Nosy Borah (formerly the He Sainte Marie) off the east coast; the 
population is mixed though originally Indonesian, later influenced by Arabs and 
pirates. The island was ceded to the French in 1750 and became the first enduring 
French settlement. 



II.4. AGRICULTURE 

Agriculture is by far the most important activity on the island and the great majority (over 
80%) of the population are either pastoralists or farmers. Agriculture supplies most of the raw 
materials for industry, ca 80% of revenues, and 34% of G.N. P. In common with almost all 
other countries, however, the proportion of the population engaged in agriculture is declining - 
FAO quote estimates of 89.4% in 1970, 83.7% in 1980 and 80.1% in 1984. 

Cultivation is the dominant form of agriculture and provides most of the national diet as well 
as the bulk of exports; stockraising is the chief activity in parts of the southern and western 
regions but is of more limited economic importance because of its low productivity. 



■14- 



Human geography 

In 1972 it was estimated that 56.6% of crop production and 73% of livestock production was 
for subsistence consumption, with some 80% of the nation's farmers engaged primarily in 
subsistence production, though most sold some portion of their crop for cash. 

Agriculture on the island is characterized by a high diversity of production, made possible by 
the wide range of climatic and edaphic conditions. It is however beset with problems at 
present, brought about partly by this very diversity of conditions, and also by the 
fragmentation and dispersal of arable land, the distance between producing areas and internal 
and external markets, low monetary return to farmers, lack of modern techniques and capital, 
and the vagaries of a tropical climate, prone to cyclones and drought. 

Rural communications are a severe problem, with only some 6300 km of road motorable 
throughout the year out of a total of around 40 000 km, much of which is dry-season tracks 
and trails. Food output has increased to some extent since the mid 1970s, but has been 
outstripped by the population increase, and with general stagnation of agricultural production 
in the 1970s, farmers have given more attention to feeding their families than to export crops 
or agro-industry. Rapid population growth, particularly in the Hauts Plateaux region, coupled 
with lack of adequate fertilizers has led to unsustainable crop rotation and soil degradation. 
Irrigation networks have not been maintained. Information is lacking and responsibility for 
agriculture is dispersed between various ministries and para-statal organizations. 

The 1978-79 development plan allocated 55 400 million F.M.G. to agricultural development. 
This had limited success for a variety of reasons, including delay in redistribution of 
requisitioned land, insecurity of rural regions (increase in cattle thefts), failure to eradicate 
share-cropping, and the relatively disappointing performance of the rural Fokonolona 
co-operative societies as vehicles for participation in and promotion of rural revival. Climatic 
conditions were also particularly difficult in the early 1980s, with four cyclones in January 
1982, following a long period of drought which had itself depressed agricultural production, 
and a further four cyclones in early 1984 which destroyed an estimated 40 000 ha of 
rice-fields, mainly in the provinces of Toliara and Fianarantsoa. 

A document on agricultural policy circulated in February 1983 outlined many of the prevailing 
difficulties and proposed solutions, emphasising particularly the ironing out of structural and 
organizational problems and aiming in the short term at the rehabilitation of the existing farm 
economy. Agrarian reform, an essentially political issue complicated by centuries of tradition, 
will aim at settling under-used land, ensuring a minimum of viability on small farms (average 
holding is 1-1.5 ha) and usefully channelling internal emigration. Mechanization was to take 
second place to the encouragement of artisanal techniques and the better use of traditional 
hand-tools. 

Consequent on this, a three-year plan for the agricultural sector announced in June 1984 laid 
most emphasis on restoring and developing the island's rice production with the aim of 
restoring self-sufficiency in rice by 1988. Other areas emphasised were livestock, 
tree-planting and fisheries. 



Land use and farm structure 

In 1972, some 5 000 000 ha or 9% of the total land area was considered to be suitable for 
cultivation without extensive reclamation measures, though other estimates put the extent of 
arable land at as much as 15% of the land area. FAO estimates for 1982 gave some 
3 011 000 ha as under cultivation or fallow, a small increase over a 1968 estimate of 
3 000 000 ha. 

In 1965 land use (of cultivable land) was estimated as: 35% fallow; 25% under irrigated crops 
(mainly rice, also sugar and cotton); 10% under tree crops (e.g. coffee, cloves); 30% planted to 
dryland crops (e.g. mountain-rice, cassava, maize, groundnuts, sisal and tobacco). 



15- 



An environmental profile of Madagascar 

Trees were grown primarily in the east and north, irrigated crops in the Hauts Plateaux and 
river basins of the west; dryland crops were more evenly distributed, accounting for around 
37% of cultivated areas in the east, 46% in the Hauts Plateaux, 53% in the north and 58% in the 
west. 

In 1971 there were estimated to be around 940 000 farms on the island. The great majority of 
these were small private (family) holdings, the average size having been estimated at from 1 ha, 
excluding forest and right of way, to 1.7 ha in all. This size does not appear to vary much 
from region to region. Only 3% of holdings exceed 4 ha. 

Most family holdings were divided into a number of separate fields or plots (e.g. rice paddy, 
kitchen garden and field for dryland crops). Pastureland is usually communal. 



Crop Production 

Crop production, derived from FAO figures, is given in Table 3. Crops grown principally as 
cash crops include: coffee, cloves, vanilla, sugar, groundnuts, cotton, soya, coconut (for copra), 
sisal, tobacco, and cocoa; the remainder are apparently mainly grown for subsistence 
consumption.. Of cash crops, coffee accounted for 35% (in value) of all exports in 1981, cloves 
23% and vanilla 8.8%. 

a. Rice is the single most important crop and is the dietary mainstay of the Madagascan 
people. Average yearly consumption per head was put at ca 135 kg in 1972 (thus ranking as 
the world's fifth largest per capita consumer). Rice was originally a prestige food, its use has 
now spread throughout the island, replacing in most areas the traditional diet of cassava and 
other tubers. Only the people of the south (e.g. Antanosy, Antandroy) still feed mainly on 
these, and in 1972 it was noted that rice was increasing in popularity even here. 

Rice is grown on around 1 million ha and accounts for 40 to 50% of the total annual value of 
the island's agricultural produce. Annual production up to 1982 (when severe flooding had a 
disastrous effect) was more or less static at just over 2 million tonnes despite the considerable 
governmental efforts to improve rice production in the 'fight for rice'; moreover a decreasing 
proportion of the crop was reaching the open market: normally some 88% is retained by the 
growers for domestic consumption, but this had increased to perhaps 96%, as a result of 
increasing population pressure and the deterioration of both irrigation systems and trade and 
transportation networks in most areas. A 27% price rise imposed by the government in 1982 
appears to have helped alleviate this. Up to 1972 Madagascar was a net exporter of rice, by 
1982 the country had to import 15% of its needs. The 1981 import of 170 000 tonnes absorbed 
8.5% of the country's revenues. 

In 1965 an estimated 85% of riceland was in irrigated paddies ('tanim-bary'), the remaining 
15% was rain fed ('horaka') or cultivated by the 'tavy' method of slash-and-burn. About half 
of the country's irrigated rice production was in the Hauts Plateaux including the Alaotra 
basin, this last area is the site of a massive agricultural development project for rice cultivation. 

In the early 1960s four sub-prefectures produced a substantial amount of rice surplus to their 
needs. Three of these were in the Hauts Plateaux: two in the Alaotra basin and one on the 
Antananarivo plain. The fourth was on the swampy Marovoay plain along the Betsiboka River 
on the western coast, an area where a modern agricultural station and a large-scale French 
private firm had been promoting the cultivation of high quality rice varieties for export to 
France and Mauritius. At this time the densely populated Imerina and Betsileo regions of the 
highlands tended to be in overall balance of rice production and consumption, though by the 
early 1970s the situation had already started to deteriorate. Other areas roughly in balance 
were largely those with a relatively low population density or a strong production of export 
crops, such as the north-central region and the sedimentary riverain areas of the western coast. 

Yields are generally very low compared with other countries - the average for the country in 
1970 was a mere 1.7 tonnes per ha, with rain-fed rice producing around 1.2 tonnes and tavy 

-16- 



Human geography 



TABLE 3. PRINCIPAL CROPS (000 tonnes) 
Year 1974-76 1982 1983 



1984 



Rice 


2009 


1970 


2147 


2132 


Cassava 


1321 


1898 


1992 


2047 


Sugar cane 


1412 


1409 


1621 


1660 


Sweet Potatoes 


340 


356 


463 


463 


Potatoes 


124 


201 


253 


264 


Bananas 


423 


281 


286 


224 


Mangoes 


194 


160 F 


160 F 


170 F 


Maize 


125 


113 


132 


141 


Taro 


82 


78 


85 


93 


Coconuts 


32 


80 


80 


82 


Oranges 


77 


80 


81 


81 


Green coffee 


81 


81 


81 


81 


Pulses 


71 


55 


58 


57 


Pineapple 


56 


50 


50 


50 


Beans (dry) 


62 


41 


45 


44 F 


Ground nuts (+ shell) 


45 


32 


31 


32 


Cottonseed 


22 


16 


16 


20 


Sisal 


29 


18 


19 


20 


Avocados 


18 


13 F 


13 F 


14 F 


Cotton (lint) 


11 


10 


10 


12 


Copra 


4 


11 


10 


10 


Cocoa 


4 


6 


6 


6F 


Palm kernels 


+ 


5 


5 


5 


Cashew 


3 


4 


4 


4 


Tobacco 


4 


3 


2 


3 


Palm oil 


2F 


3 F 


3F 


3 F 


Castor beans 


1 


1 F 


1 F 


1 F 



Figures for cloves, vanilla and pepper have not been located for these years. 
F = FAO estimates 

Source: FAO Production Yearbook 1984. 



-17- 



An environmental profile of Madagascar 

method 1.5 tonnes. Even irrigated paddies produced results far below those obtained elsewhere 
- in the Antananarivo district in 1970, the average yield was 4.2 tonnes per ha, though some 
producers were obtaining 5-7 tonnes per ha in fields of 1 to 2 has, while in the large-scale 
mechanized Alaotra project up to 7.4 tonnes per ha had been obtained by the mid-1960s. This 
compares with Japanese yields at that time of 50 to 60 tonnes per ha on 1 to 2 ha fields. 

Attempts to increase rice production have been a major thrust of the government's agricultural 
policy for many years and have centred on both increasing production (through increasing 
yields and the area under cultivation) and improving net distribution, especially to town 
dwellers; these have however met with only limited success. 

Official measures include forming state companies (SINPA) to collect rice for milling and 
marketing (1973), issuing ration cards for rice and controlling sale and price to consumers 
(1975), inducing change from rice to wheat in diet (1976), creating more farmer's co-operatives 
(1977), improving irrigation over 20 000 ha by building 1500 small dams (1978), decreasing 
consumption with one riceless day per week (1979-1980) and increasing price to paddy 
producers by 17% (1981), and again by 27% in 1982. 

The effectiveness of these measures has been variable, though in general the campaign has had 
limited success. This has been ascribed to a number of causes, including drought, especially in 
the south, problems in the running of SINPA, and changing food habits of the Malagasy, with 
rice becoming ever more popular. Efforts to eliminate middle-men and usurers have 
reportedly been relatively successful, though the effects of this do not appear to have been 
completely beneficial - in large measure the state companies which have replaced them 
(SINPA) have run into serious financial and managerial problems. Although farmers may have 
previously been exploited by the traditional middlemen, these people also brought goods into 
the villages, maintaining internal trade circuits. There are now virtually no consumer goods, 
and incentive to sell produce and increase output beyond the needs of the producer's family is 
correspondingly low. 

From 1975 to around 1982, only 25 000 ha of new rice fields were under cultivation, the goal 
of the government for one million new hectares between 1978 and 2040 was already falling 
behind. However the 1983 and 1984 harvests were considerably higher than the 1982, being 
2147 and 2132 million tonnes respectively, compared with 1723 million tonnes in 1982; 
improvements such as the construction of dozens of small dams each year are thus having an 
effect. Increasing yield has been gained by increasing the area under cultivation rather than 
yield per hectare, which decreased from 1983 to 1984 and is still below that achieved in the 
period 1974-76. A Swiss backed campaign against pests (grain borers) has begun in the 
important rice-growing area of Lake Alaotra which could increase yields there by about 500 
kilos per ha. 

b. Cassava (manioc). In 1972 this was stated to be the second food crop after rice in terms of 
area planted and probably in quantity consumed, though it rated low in consumer preference. 
FAO production figures for 1980 note an unofficial yield estimate of 1 450 000 tonnes, placing 
it third after rice and sugar cane. This represents around a fourfold increase over estimated 
total production in 1962 of 327 000 tonnes, though this may be accounted for by a change in 
type of estimate i.e. from consumption to production (see below). However Thompson and 
Adloff quoted a figure of 800 000 tonnes harvested in 1961 from 202 600 ha. 

In the 1960s it was grown in every part of the island except the interior grazing lands of the 
west and the uncultivable mountain ranges of the east. Production was greatest in the far 
south, where it was a staple of subsistence consumption, and around Lake Alaotra and the 
Sambirano River, where it was grown for industrial processing into tapioca, starch and flour at 
the chief processing centres in the provinces of Antananarivo, Mahajanga and Toamasina. 
Exports were declining and not considered to have much potential, and were apparently 
insignificant by 1980. 



Human geography 

In 1962, some 100 000 tonnes out of total production of 327 000 tonnes were fed to livestock, 
especially in the Hauts Plateaux, where it was often used as fodder for penned cattle. 
Elsewhere it was likely to be used primarily as a reserve against famine, so that potential 
resources in cassava in an average year were thought to be about double the amount actually 
consumed for subsistence or marketed. Often plots would be left unharvested as a sort of 
domestic larder, either because commercial demand and price were too low in the locality or 
because subsistence consumption was directed by preference to rice and vegetables - often for 
both reasons. 

In 1972 it was stated that no increase in the area of cassava was recommended, instead better 
utilization of areas already planted was advocated. 

c. Other tubers. Sweet potatoes, potatoes and taro are produced in large quantities (see Table 
3); the increase in production of these, and maize, has partially compensated for the short-fall 
in rice production, although rice remains the greatly preferred foodstuff. Wild roots and 
tubers are also harvested, although no reliable figures for consumption are available. 

d. Coffee is the single most important Madagascan export and is grown along the east coast and 
in the north-west (in the lower Sambirano region and on Nosy Be). In 1983 it was reported to 
cover an area of around 220 000 ha along the east coast and its production is said to involve 
about 25% of the island's population. Production has been relatively stable at around 
80 000 tonnes per annum, though is noted to have peaked in 1979 at 81 000 tonnes and to have 
declined since. Most of the bushes are now well past their best and the government launched a 
scheme 'Operation Cafe arabica' in 1979 to replace them, improve cultivation techniques and 
increase the price paid to producers; although this has resulted in the planting of some 425 000 
new coffee bushes in 1981 and an increase in price to the growers of around 50% since 1979, it 
is still said to be behind schedule. Under IMF terms the government is currently committed to 
exporting 60 000 tonnes of coffee a year. Collection from outlying areas has been hampered 
by the deterioration of the secondary road network. 

e. Vanilla is grown mainly in the north-east, also in the north-west. Thompson and Adloff 
(1965) quote a figure of 5000 ha planted to the crop, with 4700 ha of this in the region of 
Antseranana. Madagascan vanilla accounts for 90% of the world's sales of that commodity, in 
1983 apparently all to USA; however it faces strong competition from synthetic subsitutes. 
Production fell during the 1970s partly because of poor plantation maintenance, from around 
8000 to 2000 tonnes per annum. Prices to the producer were more than doubled between 1979 
and 1982 and output was reportedly increasing again. 

f. Cloves are reportedly grown almost exclusively on the eastern coastal plain, on Nosy Borah 
and around Fenoarivo Atsinanana; in 1965 the crop was said to cover around 35 000 ha. Clove 
production follows a 3-4 year cycle which was reportedly at a low in 1983. Production 
decreased from 11 000 tonnes in 1979 to 8000 in 1980 (presumably this could be part of natural 
cycle), as did the Malagasy share in a joint marketing venture with Tanzania, started in 1977. 
All exports are apparently to Malaysia. However it was noted that the area under cultivation 
had been steadily increasing and producer prices raised, which could give good results in 
1985/1986. 

g. Sugar is grown on four government estates and many small-holdings; production in 1981 was 
estimated at 1.4 million tonnes. However yields from the small-holdings are low and mostly go 
into rum, with production of refined sugar having reportedly declined. A few thousand tonnes 
are exported. 

h. Cotton is grown for local processing; in 1967 it was noted that important areas of cotton 
existed in the region of Toliara in the deltas of the Fiheranana and Mangoky Rivers and 
around Ankazoabo. It was hit by drought in 1980, with yields of seed cotton dropping from 
around 35 000 tonnes in 1979 to 25 000 tonnes in 1980; by 1983 production had recovered to 
around 30 000 tonnes and producer prices have doubled over the past few years. 



•19- 



An environmental profile of Madagascar 

i. Sisal is grown principally in the arid regions of the south and south-west (together producing 
four fifths of the country's total), but also around Antseranana and on the lower Betsiboka. 
Areas conceded to sisal companies in 1960 covered 25 000 ha of which 16 000 ha had been 
planted to the crop. Production of sisal in 1965 was given as 24 300 tonnes while estimates for 
1979-80 were 22 000 tonnes per annum; production of this long-established crop has thus 
remained relatively stable. It was noted in 1967 that international competition made export 
difficult. 

j. Oil crops. Groundnut production has decreased, but the government was reported in 1983 as 
encouraging soya bean planting (on 70 000 ha) and copra from coconuts (3000 ha) and 
groundnuts (60 000 ha) and had plans to build another oil-mill. 

k. Tobacco. Production has remained relatively stable and stood at around 4000 tonnes in 1981. 



Livestock 

Estimates for livestock numbers and products are given in Tables 4 and 5. 

a. Cattle are by far the most important form of livestock and traditionally there is one head of 
cattle per person in Madagascar. In 1981 the registered herd numbered 7.3 million; estimates 
of the true number vary widely, though it is almost certainly higher than this (as reflected in 
FAO estimates quoted above). The principal cattle producing areas are the provinces of 
Toliara and Mahajanga in the west and the prefectures of Alaotra, in the Hauts Plateaux, and 
Vohimarina in the north-east. These areas in total hold some 2/3 of the national herd, and in 
1972 had a reported surplus production of 11-13%. Cattle tend to be valued more in social 
than in economic terms and official slaughtering was only 2.5% of the herd in 1981. In 
principle, cattle in the traditional pasturage areas of the south and west are only slaughtered on 
ceremonial occasions; however, such occasions are frequent enough that meat consumption is 
high - in 1972 per capita consumption of beef in such areas reportedly averaged 66 pounds 
(30 kg) each year. 

Most of the Hauts Plateaux and eastern region constitutes an area of net cattle consumption, 
with 8-12% (in 1972) of the herd being imported from cattle producing regions. Some meat is 
also exported. Overall, however, 75% of total output (in 1972) was consumed as subsistence, 
with only 25% entering the money exchange economy. Money from sales tends to go into 
replacing the herds; the livestock sector is thus something of a closed circle economically. In 
1982 less than 20% of the current export quota was being met and a recent study showed that 
the country may need to import by 1985. Supply was not a problem but commercial networks 
- rounding-up, slaughtering, veterinary care and transport - posed organizational difficulties 
which raised costs above prices. 

In 1972, cattle taxes were abolished as an encouragement to stock-raising, yet the need to pay 
such taxes had obliged farmers to sell some cattle and thereby kept something of a monetary 
circuit going. Government policy is now tending towards the suppression of agricultural 
subsidies. Permits to move cattle across country may also soon require payment. Vaccination 
has been free, but may now be charged to farmers, although in 1982 the World Bank, FAO and 
the Madagascan Government began joint financing of an immunization campaign and other 
measures in Mahajanga province to improve stock-rearing and increase the income of the 
estimated 120 000 herders in the province. The country is looking for export markets and 
hopes to sell meat this year to the EEC, North Africa and within the Indian Ocean. 

Cattle are generally free-roaming and subject to minimal husbandry; the burning off of pasture 
to provide new growth for grazing during the start of the dry season is probably the major 
cause of deforestation in western regions (see Part III. 4.). 

b. Pigs can be raised in all parts of the island; however, Thompson and Adloff noted in 1965 
that commercial hog-raising was widespread only in the Hauts Plateaux region, one of the 
principal reasons for this being that pigs and pork were 'fady' (taboo) for many of the coastal 



-20- 



Human geography 



Year 



TABLE 4. LIVESTOCK ('000 HEAD) 
1974-76 1982 1983 



1984 



Cattle 


8543 


10 281 


10 322 


10 400 F 


Goats 


1308 


1730 


1750 F 


1800 F 


Pigs 


607 


1240 


1300 F 


1350 F 


Sheep 


632 


740 


630 F 


700 F 


Chickens 


13 000 


18 000 


18 000 


18 000 


Ducks 


2000 


4000 F 


5000 F 


5000 F 


Turkeys 


1000 


3000 F 


3000 F 


3000 F 


F = FAO estimates 











TABLE 5. LIVESTOCK PRODUCTS (TONNES) 1 

Year 1974-76 1982 1983 1984 



Beef and veal 


108 


135 


136 


138 


Poultry meat 


36 


63 


66 


70 


Pigmeat 


23 


31 


32 


34 


Goat meat 


3 


9 


9 


9 


Mutton and lamb 


2 


2 


2 


2 


Milk (whole, fresh) 


30 


39 


41 


43 


Hen eggs# 


9808 


13967 


14164 


14360 


Eggs (excl. hens)# 


3012 


4320 


4500 


4680 


Honey 


11 


4 F 


4 


4 


Silk (raw and waste) 


15 


15 F 


15 


15 



1 All figures for 1982-84 are FAO estimates 
# Number of eggs ('000). 

Source: FAO Production Yearbook 1984. 



-21- 



An environmental profile of Madagascar 

tribes, including any with Arabic influence. FAO estimate that pigmeat production increased 
by just under 50% from 1974 to 1984 (Table 5). 

c. Sheep. In the early 1960s the only important area of sheep production was in the Androy 
region in the extreme south-west, although other areas were climatically suitable and the Hauts 
Plateaux region had previously supported significant numbers of sheep. As with pigs, sheep 
were considered 'fady' by several of the coastal tribes. According to FAO estimates (see Table 
4), the number of sheep decreased in the period 1982-84, although the 1984 estimate showed a 
10% increase over the 1974-76 population. 

d. Goats increased in number from 1974 to 1984 at approximately the same rate as pigs. 
Thompson and Adloff (1965) noted that these were found very largely on the west coast and, 
like sheep, mostly in the extreme south-west, despite large areas of the rest of the island being 
climatically suitable for rearing them. 

e. Poultry, especially chickens, are ubiquitous and provide an important source of protein, both 
as eggs and as meat. 



REFERENCES 

Anon. (1983). Madagascar - the 'taxi-brousse' changes gear. The Courier 80: 7-26. 

Battistini, R. and Verin, P. (1972). Man and the environment in Madagascar. In: 

Battistini, R. and Richard-Vindard, G. (Eds), Biogeography and ecology in 

Madagascar. Monographiae biologicae 21. Junk, the Hague. 
FAO (1985). FAO Production Yearbook 1984. FAO Statistics Series No. 61, Vol. 38. Food 

and Agriculture Organisation of the United Nations, Rome. 
Guilcher, A. and Battistini, R. (1967). Madagascar - geographie regionale. 'Les cours 

de Sorbonne', Centre de documentation universitaire, Paris. 
Jolly, A. (1980) A world like our own; man and nature in Madagascar. Yale University 

Press, New Haven and London. 
Nelson, H.D., Dobert, M., McDonald, G.C., McLaughlin, J., Marvin, B. and Moeller, P.W. 

(1973). Area handbook for the Malagasy Republic. U.S. Govt. Printing Office, 

Washington, D.C. 
Paxton, J. (1985). The Statesman's Year-book 1985-1986. Macmillan, London. 
Thompson, V. (1986). Madagascar. In: Africa south of the Sahara. Europa Press. 
Thompson, V. and Adloff, R. (1965). The Malagasy Republic. Stanford University Press, 

Stanford. 



-22- 



PART III. FLORA, VEGETATION AND FOREST COVER 

III.l. FLORA 

Diversity 

i. Estimates of the size of Madagascar's flora vary between 7370 and 12 000 species. The lower 
figure is that of Perrier de la Bathie (1936) and includes Pteridophytes; it is almost certainly 
too low. Largely due to research for the Flore de Madagascar et des Comores (Humbert, 
1936-), the flora is much better known now, and more recent estimates give the figure as much 
higher: 8200 species (Leroy, 1978; Raven, 1985); 8500 vascular plants known (White, 1983); 
10-12 000 species, (though this may include Pteridophytes) (Guillaumet and Mangenot, 1975); 
10 000 Angiosperms (Rauh, 1979), also quoted in White (1983); 12 000 species (Guillaumet, 
1984). Greater credence is given to the lower of these figures since they seem to be the only 
ones based on the known flora rather than on estimates. 

Lebrun (1960) gave Madagascar an area-richness index of 5.4, which he based on the figure of 
7800 from Humbert (1959). Even so, this high area-richness figure is close to that of the 
south-west cape of Africa with an area-richness index of 5.6. The latter area has been 
described as one of the richest areas botanically in the world. If the true number of species in 
Madagascar is 10-12 000, then the area-richness index should be nearer 8, which could perhaps 
make it unequalled (Guillaumet and Mangenot, 1975). 

Figures for the numbers of genera and families in the flora show rather less variation: Perrier 
de la Bathie (1936) gives 1289 genera and 191 families; Guillaumet (1984) gives 1600 genera in 
180 families; Leroy (1978) gives over 160 families; White (1983) gives 1200 genera. 

ii. The distribution of the flora between the two main regions of the island is as follows, based 
on the figures of Perrier de la Bathie (1936) and Humbert (1959), and quoted in Koechlin 
(1972): in the Eastern Region there are 500 genera and 5500 species; White (1983) gives 1000 
genera and 6100 species in this region; in the Western Region there are 200 genera and 1800 
species; White (1983) gives 700 genera and 2400 species in this region. 600 species are common 
to both regions. 605 species of the total are introduced and naturalized, and 945 species are 
native but non-endemic; the rest are endemic. 

iii. The biology of the introduced species is strikingly different from that of the native species 
(Dejardin et a/.., 1973; Leroy, 1978): 53% of naturalized species are annuals, as against only 3% 
of the indigenous flora. Remarkably, 83% of the introduced flora is herbaceous, and has 
largely not penetrated the primary forest, but keeps to the secondary, man-modified 
formations. Of the indigenous species, more than 80% are woodland plants. 



Endemism 

i. Endemic families. Out of 191 families, White (1983) identifies eight as endemic: 
Asteropeiaceae (1 genus, 5-6 species); Didiereaceae (4 genera, 11 species); Didymelaceae 
(1 genus, 2»species); Diegodendraceae (monotypic); Geosiridaceae (monotypic); Humbertiaceae 
(monotypic); Sphaerosepalaceae (Rhopalocarpaceae) (2 genera, 14 species); Sarcolaenaceae 
(Chlaenaceae) (10 genera, 35 species). Guillaumet (1984) gives "6 or 7" endemic families. 
Perrier de la Bathie originally listed three as endemic: Rhopalocarpaceae, Chlaenaceae and 
Didiereaceae. 

The endemic Didiereaceae is ecologically the most important family on the island, forming the 
most striking feature of the spiny desert in the south and south-west. Certain of the species 
resemble the American Fouquieriaceae in habit, and the family is reputed to have strong 
affinities with the Cactaceae (Rauh and Reznik, 1961). 

ii. Endemic genera. Of the 1289 genera to which Perrier de la Bathie (1936) refers, 238 (20%) 
are endemic . These figures are also quoted in White (1983). 



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An environmental profile of Madagascar 

iii. Endemic species. Out of the whole flora, Perrier de la Bathie gives a figure of 5820 species 
(86%) as strictly endemic. Humbert (1959) gives the figure 6400 or 81% of his known 7800 
species. Raven (in press) estimates that 4500-5000 species (55-61%) are endemic. Guillaumet 
(1984) says 85% of his 12 000 species are endemic. 

Endemism is most marked in woodland and forest formations where an estimated 89% of 
species are endemic (Perrier de la Bathie, 1936). Other figures are: rocky outcrops, 82% 
specific endemism; marsh, 56%; littoral formations, 21%. 



trees, 94%; shrubs, 



Percentages of specific endemism are given for different biological types: 
94%; perennial grasses, 85%; annual grasses, 58%. 

iv. Area based endemism. Area based endemism (generic/specific) is as follows (figures from 
Perrier de la Bathie, 1936): South Domain: 48% and 95%; West Domain: 41% and 90%; East 
Domain: 37% and 90%; Sambirano Domain: 23% and 89%; Centre Domain: 21% and 89%. At 
the regional level, endemism is more pronounced in the Western Region, with 38% generic and 
89% specific endemism; White (1983) gives 20% generic and 79% specific endemism. In the 
Eastern Region Perrier de la Bathie (1936) gives 22% generic and 82% specific endemism; 
White (1983) gives 16% generic and 79% specific endemism. Of the species which are common 
to both, there is 9% generic and 47% specific endemism. 



Affinities of the flora 

On the basis of his species counts, and his knowledge of the Madagascan flora and that of the 
rest of the world, Perrier de la Bathie (1936) quantified the relationships between the 
Madagascan and other floras of the world . These figures are widely quoted (Dejardin et al., 
1973; Koechlin, 1972; Koechlin et al., 1974; Leroy, 1978) although there seems to have been 
some guesswork in their origin. 

Around 27% of the Madagascan flora is estimated to have African affinities; the pantropical 
element is around 42%; the Oriental element 7%; the austral element accounts for 3%, and the 
recent element whose transport is due to long distance dispersal 15%. The remaining 6% 
represents the true endemic element. In addition to these, Dejardin et al. (1973) recognise the 
existence of a neotropical element, and also a paleotropical one. They also unite Perrier de la 
Bathie's recent and exotic elements. 

Exactly what these figures represent is unclear, but phytogeographers often quantify foreign 
elements as the number of species in the flora belonging to genera which also occur in the 
relevant foreign countries or areas. 

i. All cosmopolitan and pantropical families of both Pteridophytes and Angiosperms are 
represented in Madagascar (Dejardin et al.. 1973), and account for a large proportion of the 
flora. In contrast, Gymnosperms are represented only by the genus Podocarpus. Most of the 
species are ruderals, or weedy plants of grassland, borders and clearings, or are hydrophytes. 

ii. The African element is related to the Sudano-Zambesian flora of eastern and southern 
Africa rather than the Guineo-Congolian flora of the west; notable examples include the 
genus Coffea, which is represented in Africa by 15-20 species, and in Madagascar by about 50 
species (Guillaumet and Mangenot, 1975). Among bryoflora, the family Rutenbergiaceae is 
striking, being found in Madagascar and the Mascarenes, and also on the old crystalline 
mountains of Tanzania (Pocs, 1975). Leroy summarises the African element in some detail, 
dividing it into seven broad types, and citing numerous examples. 

iii. The Oriental element is much less important than the African element, accounting for 7% 
of the flora including (following Dejardin et al.,) some seven families and two subfamilies 
(Dejardin et al., 1973). Notable generic examples (out of some 50 cited) include Nepenthes 
and Pandanus. The former has its centre of diversity in Borneo and Sumatra and eastern limit 



-24- 



Flora, vegetation and forest cover 

in New Caledonia; there is one endemic species in Sri Lanka, one in the Seychelles and two in 
Madagascar, (N. madagascariensis plus one undescribed from Cape Masoala) (Dejardin et al., 
1973). Pandanus is distributed in all the floras of the tropical Old World, Australia and 
Oceania, but is completely absent from America. About 700 species are described with some 
550 in Asia and Oceania. 76 species occur in Madagascar, and all but two are endemic. A 
further 30 species occur in the Mascarenes, the Comoros and the Seychelles, and only 22 in 
Africa (all but one in the east and centre). 

iv. The Southern element appears somewhat less well defined; Perrier de la Bathie considered it 
accounted for around 3% of the flora and included in it a few species in common with South 
Africa, several endemic species related to plants in South Africa, and 77 genera in common 
with South Africa, South America or Oceania, or relatives of genera confined to those areas. 
The most important of these are African. Dejardin et al. adopted a somewhat broader view, 
including in this all taxa which are predominantly found in the southern hemisphere, and those 
that are confined to some part of the southern hemisphere, either African or eastern. Other 
taxa which are now widespread in the northern hemisphere probably have a southern origin, 
for example the palms, which show considerable diversity in Madagascar (see Appendix 4). 
Dejardin et al. (1973) recognise 13 families and subfamilies in this element concluding that the 
southern element in the Madagascan flora probably exceeds 3%. 

v. The Palaeotropical element is defined by Dejardin et al. (1973) as the tropical taxa abrent 
from continental America but not confined to any one region of the Old World Tropics; 8 
families and 73 genera are cited in this group. 

vi. There is a small but significant element in the Madagascan flora which has affinities with 
America - the Neotropical element. These taxa show a variety of distributions from wide 
continuous ranges, like that of Sabicea (with c. 50 species in Tropical America and the West 
Indies, 85 species in continental tropical Africa, and 5 species in Madagascar), to the extremely 
discontinuous ranges shown by Ravenala (with one species in Madagascar and one in Brazil and 
Guiana) and Oplonia (with one species in Peru, 8 species in the West Indies and 5 species in 
Madagascar). Stearn (1971) cites 9 families restricted to America and Madagascar only, while 
Dejardin et al. (1973) cite 26 genera which are common to Madagascar and America, and 
present or not in Africa. 

vii. Undoubtedly, a substantial part of the Madagascan flora has been introduced relatively 
'recently' and by long distance dispersal. The exact definition of 'recent' in this context is 
unclear, since it could include long distance dispersal by way of oceanic islands any time after 
the breakup of Gondwanaland, and so overlap with most of the other floristic elements. Some 
species introduced by man (e.g. species of Pavonia, Sida, Cyperus) have already given rise to 
new endemic species or varieties (Koechlin, 1972). 

Origins of the flora 

Theories of continental drift or plate tectonics help provide explanations for many of the plant 
distributions outlined above. Raven and Axelrod (1974) review the state of knowledge to date, 
with particular reference to the origin of the Angiosperms. Where not otherwise stated, the 
original source of the following information is Raven and Axelrod (1974). 

During the Jurassic, the major landmasses existing today were joined together into two large 
continents: Laurasia, including North America, Europe and Russia; and Gondwanaland, 
including Africa and most of the rest of the southern hemisphere. Africa as we know it was 
joined to South America, and on the other coast Madagascar was nestled between Africa, India 
and Antarctica. By the upper Jurassic, about 150myBP, these supercontinents had started to 
break up into recognisable shapes, and move, by a process not yet fully understood, across the 
surface of the globe. Antarctica and Australia moved south, while South America moved west, 



-25- 



An environmental profile of Madagascar 

leaving Africa, Madagascar and India still joined together. From evidence reviewed in Axelrod 
and Raven, 1978, it is likely that Madagascar and India remained together longer than they 
remained attached to the African mainland, and that they split from Africa sometime in the 
mid-late Cretaceous (100-80myBP); India and Madagascar probably separated in the early 
Paleocene (c.65myBP). 

The exact position of Madagascar during the Jurassic has been a point of considerable 
controversy, but according to Smith (1976a, 1977b) the matter is settled with the presentation 
of palaeomagnetic evidence. This is based on the comparison of the orientation of the 
magnetism solidified into the ancient rocks of Madagascar and the eastern part of Africa. The 
two possibilities most often presented are: 1. that Madagascar was attached to the east coast of 
Somalia/Kenya/Tanzania, the so-called northern position; and 2. that Madagascar originated in 
a southern position, attached to the east coast of Mozambique, more or less opposite its present 
position. The overwhelming evidence to date, both geological and biological, suggests a 
northern Gondwanaland position for Madagascar (Embleton and McElhinny, 1975; Hilliard and 
Burtt, 1971; Smith and Hallam, 1970; Smith (1976a); Smith (1976b); Wild 1975) and that the 
island has subsequently moved south and east to its present position. 

According to Raven and Axelrod (1974), and concurred by Leroy (1978), primitive 
angiosperms originated in the western part of Gondwanaland, probably in the lower Cretaceous 
(c.l25myBP). By the end of the lower Cretaceous, before Laurasia had separated from 
Gondwanaland, these had spread north and diversified, so that by the upper Cretaceous a large 
flora was established in each of the two supercontinents: Magnoliaceae, Ranunculaceae, 
Amentiferae, etc. in Laurasia; Annonaceae, Winteraceae and Myristicaceae in Gondwanaland. 
By the early Paleocene, almost all modern families were in existence, and had spread to the 
peripheral parts of each continent. 

The spread and subsequent diversification of this Gondwanan angiosperm stock explains the 
presence in Madagascar of a large pantropical and cosmopolitan element. The very oldest 
affinities are those at the highest taxonomic level, and well distributed throughout all parts of 
Madagascar. The southern element, for example, is attributed with being one of the oldest in 
Madagascar; the taxa have marked archaic characteristics and are distributed all over the island 
(Koechlin, 1972). As the oceans opened up, the possibility of migration was reduced, and 
subsequent diversification of the separated parts raised the level of the floristic affinities. 

The American element in Madagascar, and some parts of the southern element, can be 
explained on this basis, if the original stock inhabiting the South American and African 
landmass was split when these two continents parted, and if there were sufficient changes in 
the environment of continental Africa for a proportion of this stock subsequently to become 
extinct. There is considerable evidence now that there were indeed major climatic changes in 
Africa concurrent with the Quaternary glaciations in the polar regions; particularly the 
widespread occurrence of aridity (Axelrod and Raven, 1978; Coetzee, 1964; Coetzee and van 
Zinderen Bakker, 1970; Diamond and Hamilton, 1980; Hamilton, 1981; Livingstone, 1975; 
Raven and Axelrod, 1974; Richards, 1973; Stearn, 1971). This could have caused widespread 
extinction of taxa in Africa compared with South America and Madagascar, whose climates 
appear to have remained stable for a very long time. Richards (1973) highlights the uniqueness 
of the African forests, and remarks on the paucity of their flora. 

Because of its relatively recent attachment to the African continent, Madagascar has very 
strong African links in its flora. This is borne out by the fact that most of these affinities are 
at species level. Even after the continents had split apart, there was the possibility of 
migration across water on numerous small islands probably into the upper Miocene, although 
this became less as the distances increased. The existence of dwarf hippos in both Africa and 
Madagascar suggests links up until the end of the Pliocene (Koechlin, 1972). 



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Flora, vegetation and forest cover 

III.2. VEGETATION 

The following is a summary account of the native vegetation of Madagascar. More 
comprehensive descriptions, from which this is taken, are found in Humbert (1959), Humbert 
and Couts Darne (1965), Koechlin et al. (1974), Perrier de la Bathie (1936), and White (1983). 
The terminology and delimitation of vegetation types are after (White, 1983). 

Descriptions of the secondary formations are given below in a separate section. 

Eastern Region 

The Eastern Region (or East Malagasy regional centre of endemism) covers the eastern part of 
Madagascar and includes the central highlands which run most of the length of the island. It 
extends from the highlands eastwards down to sea-level and westwards down to an altitude of 
approximately 800 m. The Sambirano Domain is separate and forms a small enclave on the 
north-west coast. 

The original vegetation of the region was almost entirely forest. Evergreen rain forest occurred 
everywhere below 800 m, and three other forest types above 800 m: moist montane forest and 
sclerophyllous montane forest occurred on the eastern slopes of the highlands, while 'Tapia' 
forest grew on the western slopes. Above 2000 m forest is replaced by montane thicket. 
Rocky outcrops above and below 2000 m support rupicolous communities. 

i. East Malagasy primary lowland rain forest. This once covered the whole area along the east 
coast up to about 800 m, the region being characterised by high rainfall (usually more than 
2000 mm, rising to 3000 mm or more) and low altitude. On windswept ridges it is replaced by 
shorter forest with a canopy of 10-15 m. 

The main canopy is at 25-30 m, but there are no large emergent trees as in most other types of 
tropical moist forest. The lower canopy is made up of small trees and large shrubs with larger 
and less coriaceous leaves than in the upper canopy. Deciduous species are absent except near 
the upper altitudinal limit. Palms are better represented than in African forests (Richards, 
1973), especially below 200 m. The dwarf palm genera Dypsis and Neophloga are important in 
the lowest layers. Bamboos, including lianoid species, are scattered throughout all layers. Tree 
ferns occur at low altitudes, but are rarer in this forest type than in submontane forest. Large 
humus-collecting epiphytic ferns grow on the trunks of trees. Epiphytes are abundant and 
increase with altitude; especially common are Orchidaceae, Melastomataceae and ferns, the last 
of which tend to grow on the stems of tree ferns. 

This type of forest is very rich in species, varying in species composition from place to place. 
Dominance by individual species or groups of species does not occur. 

ii. East Malagasy moist montane forest. This grows chiefly between 800 and 1300 m, but may 
ascend to 2000 m in sheltered places on well-watered soils; it is transitional between lowland 
rain forest and the sclerophyllous forest of higher altitudes. The main canopy is lower than in 
lowland rain forest, at 20-25 m, and the trees are often branched from near the base, so that 
the boles are rarely straight. The leaves in the canopy are more sclerophyllous; epiphytes, 
especially mosses, are more abundant. Lichens cover the trunks and branches in an almost 
continuous layer. Large lianes are abundant, especially Compositae, Rubiaceae and monocarpic 
bamboos. The herb layer is better developed and includes ferns and other species with large 
non-xeromorphic leaves. 

Moist montane forest is as rich in species as lowland rain forest. There are many species 
endemic to this formation, but others, such as Podocarpus madagascariensis, extend from sea 
level to the summits of the highest mountains. 

iii. East Malagasy sclerophyllous montane forest. This occurs between 1300 and 2300 m, but 



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An environmental profile of Madagascar 

also lower than 1300 m on shallow soils of exposed ridges. Compared with moist montane 
forest, the environment is characterised by lower temperatures with greater daily and seasonal 
variations, stronger winds, and greater variation in humidity. 

The canopy is lower than that of Malagasy moist montane forest (10-13 m) and less well 
differentiated from lower strata, since most of the trees are branched more often, and lower 
down. Structurally it is intermediate between forest and thicket, but closer to the former. The 
leaves of the canopy species are smaller and more xeromorphic. When the canopy is relatively 
open, several ericoid species, which are more characteristic of montane thicket, occur in the 
understorey. Bryophytes and lichens are more abundant than in moist montane forest, and 
many trees are covered with the lichen Usnea. The ground is covered with a thick layer of 
bryophytes and lichens. Included in it are species more usually found as epiphytes, such as 
ferns, orchids and Peperomia; most of the layer consists of pleurocarpous mosses, but there are 
also mounds of Sphagnum. This type of forest is very susceptible to fire, which can easily 
spread through the thick layer of humus. 

iv. East Malagasy 'Tapia' forest. This grows between 800 and 1600 m on the western slopes of 
the upland massif. It is much drier than forest on the eastern slopes, lying in the rain-shadow 
of higher ground further east. Temperatures are higher, and insolation is more intense, and 
because of its resulting sensitivity to fire, 'Tapia' forest has mostly been replaced by secondary 
grassland or open woodland. 

Tapia forest is similar in appearance to Mediterranean Cork Oak forest, but there are more 
species present in the canopy, which is at 10-12 m. The canopy trees have evergreen, often 
small coriaceous leaves. Although the crowns are in contact they cast little shade, so there is a 
well developed understory, composed largely of ericoid shrubs. Lianes are quite frequent, but 
small. Tree ferns are absent, and the only palm is Chrysalidocarpus decipiens which is 
confined to the wettest places. Epiphytes are rare, with only a few small ferns and species 
of Bulbophyllum. Lichens are the only epiphytes at the lowest altitudinal limits. There is no 
ground layer of bryophytes. 

The canopy is composed mostly of Uapaca bojeri, for which Tapia is the local name. It is very 
resistant to fire, and often persists as the dominant species of secondary open woodland long 
after its normal associates have disappeared. 

v. East Malagasy montane bushland and thicket. This occurs wherever conditions are suitable 
between 2000 m (or exceptionally 1800 m) and 2876 m, the highest point on the island. It is an 
extremely depauperate derivative of montane sclerophyllous forest, and is found above it. It 
consists of a single stratum of woody plants, most of which have ericoid habit with short 
twisted stems. It is never more than 6 m high, and is often impenetrable. All the species are 
evergreen, and most have ericoid, cupressoid or myrtilloid leaves. 

Lianes and vascular epiphytes are almost completely absent, except for a few small orchids, 
though epiphytic bryophytes and lichens are plentiful. There is a discontinuous ground layer 
of bryophytes and lichens on poorly drained sites. The field layer is poorly developed. 

vi. East Malagasy rupicolous shrubland. The most characteristic vegetation of large rocky 
outcrops on the African mainland is bushland and thicket. On similar outcrops in Madagascar 
the tallest plants rarely exceed 2 m, and so Malagasy rupicolous vegetation is classified as 
shrubland even though there are some floristic affinities with African rupicolous bushland. 

Rupicolous communities do occur in western Madagascar, but are most frequent in the Central 
Domain and the Domain of the High Mountains in eastern Madagascar. They also occur in the 
Oriental Domain, but the flora is poor and not well known. 

The plants are rooted in crevices or, more often, in mats of coarse shallow soils on less steep 
slopes, and are easily washed away by heavy rainfall. The soil dries out rapidly after rain and 
the plants are strongly xeromorphic as a result of strong sunlight during the day, low 
temperatures at night, and strong winds. 

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Flora, vegetation and forest cover 

Species diversity is not very high, but there is a wide range of growth forms including leaf 
succulents, stem succulents, reviviscent pteridophytes, grasses, cactoid or coralliform species, 
and spartioid species with large subterranean structures and a fugaceous rosette of leaves borne 
at ground level. 

The presence of grasses in this formation is interesting, since it is possible that they have 
spread from these habitats to dominate the grasslands which now cover so much of the island. 

Above 2000 m, the flora on rocky outcrops becomes very impoverished, although some genera 
grow to quite high altitudes, and are often represented by endemic species on each mountain. 
A few genera (for example Sedum) are confined to this high altitude rupicolous shrubland. 
There are also many bryophytes and lichens, including the fruticulose Cladonia pycnoclada. 

Western Region 

The Western Region (or West Malagasy regional centre of endemism) covers the western side of 
the island up to 800 m. From the flat plains on the west coast, which are wider than the east 
coast equivalent, the land slopes up slowly towards the east. Because the region is in the rain 
shadow of the south east monsoon, there is a dry season of seven or more months. 

The Western Region is less rich floristically than the Eastern Region, but the flora is large and 
varied. As with eastern moist forests, single species or groups of species are very rarely 
dominant. 

There are two main types of primary vegetation: dry deciduous forest and deciduous thicket 
(the so-called spiny desert). As discussed below, most of this has been destroyed, so that 
secondary grassland is the predominant feature. 

i. West Malagasy dry deciduous forest. This used to cover most of the Western Domain, where 
the mean annual rainfall varies from 500 mm in the south to 2000 mm in the north. Most of 
the wet season rain is brought from the north or west, rather than by the south-east monsoon. 
The mean annual temperature is mostly 25-27°C. 

Dry deciduous forest is less dense than most of the moister forests in the east, with a rather 
open upper canopy at 12-15 m, and with scattered emergents to 25 m. Lianes are abundant 
and the shrub layer is well developed. The soil is mostly bare except for small patches of 
Acanthaceous subshrubs which die back in the dry season. There are very few vascular 
epiphytes (a few small orchids in the wetter types), and few lichens. Ferns and palms are 
absent, as are bryophytes. The trees of the main canopy are always deciduous. Some species 
stay green for about four months, while others lose the last of their old leaves only when the 
new ones unfold, which happens in all species shortly after the rains begin. Some species of, 
for example, Adansonia, Dalbergia and Cassia, flower before the new leaves appear. Some 
herbs, for example species of Kalanchoe and Plectranthus, have large membranaceous leaves 
during the rainy season and small leaves in the dry season. 

Excluding riparian forest, there are three main types of dry deciduous forest, growing on 
different substrates: 

- On lateritic clays. These soils, derived from basalt and gneiss, support the most luxuriant 
type of dry deciduous forest. The humus layer is deeper than in the moister forests in the east. 

- On sandy soils. These are derived from Liassic, Jurassic and Cretaceous sandstones. The 
forest is similar to that on lateritic soils, but is shorter, especially on dry soils, where it is more 
like thicket. 

- On calcareous plateaux. Forests of this type are similar to those on sandy soils but lower. 
There are fewer lianes and evergreen species. Trees and shrubs with swollen stems, 



-29- 



An environmental profile of Madagascar 

e.g. Adansonia, Bathiaea. Harpagophytum, are relatively more abundant. Among the rocks, 
height falls off very quickly, the upper and lower canopies merge, and the forest turns to 
thicket; lianes and shrubs appear, and succulents and plants with swollen stems become more 
numerous. 

Dry deciduous forest also occurs in small areas in the moister parts of the Southern Domain, 
but only as a transitional form between deciduous thicket and forest; members of the 
Didiereaceae occur in the canopy. 

ii. West Malagasy deciduous thicket. This characterises the dry Southern Domain, which has 
300-500 mm rainfall per year, most falling in the summer as local heavy showers. The dry 
season lasts at least eight months, and because the rainfall is so irregular, droughts can last 
from 12-18 months. The soil substrate is usually shallow and stony. 

The height and density of the thicket vary enormously in relation to rainfall and soil moisture. 
The shorter, more open types are mostly confined to rocky situations. Most undamaged stands 
are impenetrable or nearly so. At one extreme there is a transition to dry deciduous forest, and 
at the other, on shallow soils, the canopy is less than 2 m tall. 

The thickets are commonly 3-6 m tall, and they may have a very discontinuous stratum of 
emergent trees, exceptionally reaching 8-10 m tall. Otherwise there is no stratification, and 
the thicket consists of a complex mixture of plants of different sizes. Lianes are numerous and 
rather small. The ground flora is sparse. Many species are spinous. Physiognomically, the 
most distinctive elements are the Didiereaceae and arborescent Euphorbias; these are usually 
present in, and almost restricted to, this vegetation type. 

Didiereaceae is a small endemic family of bushy or arborescent pachycauls of distinctive 
ascending branched habit, with fascicles of very small leaves scattered along the main stems. It 
consists of four genera and twelve species: two species of Didierea, D. madagascariensis and D. 
trollii; the genus Alluaudia with six species; two species of Alluaudiopsis, and Decaryia 
madagascariensis, in a monotypic genus. Some reach 8 m or more, though most are lower. 

The photosynthetic organs are very variable in behaviour and structure. In some species, large 
leaves appear suddenly after heavy rainfall, and are shed equally suddenly. In other species, 
more fugacious leaves are produced irregularly. Many shrubs have narrow greyish leaves, 
which persist longer so there may still be leaves on some shoots while shoots in another part of 
the plant have a crop of new leaves. A few rare shrubs are evergreen. Many species have 
green photosynthetic stems, and may or may not also produce fugacious leaves. In some of 
these, for example some cactiform Euphorbs, and species of Cissus and Asclepiadaceae, the 
photosynthetic stems themselves are caducous. Several taller species have distended water 
storage stems of characteristic bottle-like appearance. 

Mangroves are discussed in Part IV. 



Secondary Formations 

i. East Malagasy secondary lowland rain forest. Although the area of degraded formations in 
the Eastern Region is very large, little of it is secondary forest (which is known locally as 
Savoka). There are two main reasons for this: firstly, Madagascar has very few indigenous 
secondary forest species, and these are much less vigorous than the corresponding species in 
tropical Africa. Secondly, the indigenous secondary forest species are unable to compete 
successfully with smaller plants which are not indigenous, or with the introduced tree 
species Psidium cattleianum and P. guajava which are more invasive than the indigenous 
secondary forest species. 

The most important indigenous secondary forest tree is Ravenala madagascariensis (Musaceae), 
the Travellers Tree, which is widespread in eastern Madagascar to 800 m. Other indigenous 



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Flora, vegetation and forest cover 

secondary forest species include Harungana madagascariensis, Psiadia altissima, species 
of Canarium, Croton, Dombeya and Macaranga, and the bamboo Ochlandra capitata. 

The ease with which secondary forest can become established is largely dependent on the 
nature of the soil. Forest regeneration is favoured by porous soils of good structure and rich in 
decomposing minerals. Compact ferallitic soils formed from impoverished parent material do 
not support secondary forest, and destroyed forest is replaced by grassland with Ravenala, 
although the Ravenala is progressively eliminated by fire. Coastal forest on leached sand dunes 
is replaced by a totally different type of secondary forest dominated by Philippia. This is 
highly combustible and degrades rapidly to grassland. 

ii. East Malagasy secondary grassland. This formation covers enormous areas, particularly in 
the Central Domain. Several variants have been recognised: 

- In the coastal region, a succession of grass associations follow after the destruction of forest 
regrowth by fire. On certain soils this type of grassland can revert to forest if fire is excluded. 

- The grassland of the 'Tanety', the region of hills between 1200 and 1500 m, has widely 
spaced tufts of low ground cover with hard infertile soil in between each. Throughout most of 
the area, one species of grass is dominant: Aristida rufescens. 

- The grassland of the 'Tampoketsa', the plateaux to the north and north-east of Tananarive, 
is very uniform floristically and dominated by very few species; the total flora is only 34 
species. 

- The grassland on the western slopes occurs between 800 m and 1600 m on ground which 
formerly supported Tapia forest, and is intermediate in stature between the short grasslands of 
the plateaux and the taller grasslands of the Western Region. Again, this formation is 
floristically very poor and homogeneous. 

- The grassland on the mountain slopes above 2000 m replaces most of the montane bushland 
and thicket which used to occupy much of the Eastern Region; it seems that this formation is 
fairly recent. It is maintained by regular grazing and burning. 

Hi. West Malagasy grassland. More than 80% of the surface of the western Malagasy Region is 
covered with secondary grassland or wooded grassland, and this is burnt each year. The 
dominant species are taller than those of the eastern Malagasy grassland, with broader flat, 
ribbon-like leaves with less sclerenchyma. 

The presence of trees and bushes distinguishes the secondary grasslands in the west from those 
in the east. Tall trees are rare, and are either relics from forest or, like the palm Medemia 
nobilis, grow otherwise on hydromorphic soils. Most trees are no more than 8-12 m tall. 
Apart from their ability to coppice after fire, the trees show few adaptations to fire, and this is 
an argument in favour of the recent origin of the communities in which they live. 

The grasslands of the western region are very poor, despite their great extent and their varied 
climate and edaphic conditions. There are at the most 300 species, and more than half of these 
are transient ruderals. If those species which only grow in the shade of trees, or are confined 
to swampy places, are also excluded then only 84 heliophilous species are left which occur on 
well drained soil and can withstand annual burning. 

Most species in the western grassland are neither characteristic nor faithful, and this reflects its 
origin. Most species are introduced from other countries, or are forest species which can 
survive destruction of the forest without showing notable modifications. Of the 84 typical 
grassland species, 31 are adventive. Of the others, 42 are certainly, and 11 probably 
indigenous. Of these, 24 have originated in forest, 4 in the deciduous thicket in the south, and 
2 species of palm {Medemia nobilis and Borassus madagascariensis) are from riparian forest. 
Eighteen other species are from the dry deciduous forest. 



-31- 



An environmental profile of Madagascar 

Only 18 endemic species are confined to the grasslands. Four are phanerophytes; eight are 
hemicryptophytes. Certain other species which are confined to grassland in Madagascar also 
occur in Africa or Asia. The presence of these heliophilous endemics is evidence for the 
occurrence in Madagascar before man of small open communities which perhaps occupied 
stations least favourable to forest, such as compact soils and rock outcrops. In such situations 
the forest was probably stunted, with an open canopy, permitting the persistence of shade 
intolerant species. 



III.3. FOREST COVER AND DESTRUCTION 

There are no recent measurements of either the extent of surviving forest cover on Madagascar 
or the rate of its destruction; figures for the former are usually derived from estimates made in 
1958 from aerial photos taken mostly in 1949 (Guichon, 1960), those for the latter are taken 
from a study by Delord in 1965 concerning the eastern region. It appears that recent 
quotations for surviving forest cover are usually extrapolations from the 1958 figures using the 
estimated rate of destruction from Delord. However, there are discrepancies between various 
estimates, such as those given by Chauvet (1974) and the FAO Tropical Forest Resources 
Assessment Project (1981) although both are apparently based on the same sources; these may 
in part be a result of differing classifications and criteria for inclusion of forest types, though 
this should be clarified. 

The most important consideration, however, is that at least 70% (and probably 75%) of the land 
surface of the island no longer has significant native woody plant cover, and that this 
percentage is increasing annually. These deforested areas are mostly pseudosteppes and 
savannas with very low species diversity. The floristic composition of these and other 
vegetation types are discussed above. 

Summary approximations as given by Chauvet (1972), and derived from Guichon (1960) are: 

Eastern type 6 150 000 ha 

Western type 2 550 000 ha 

Southern type 2 900 000 ha 

Transition 900 000 ha 

Total 12 500 000 ha 

(21% of land area) 

Degraded forest 4 300 000 ha 

Overall total 16 700 000 ha 

(28% of land area) 

With the rapid rate of forest destruction (discussed in more detail below), more recent estimates 
will obviously be lower than this. 



Eastern Forests (including Sambirano Domain) 
i. Area 

FAO/UNEP (1981) quotes the following figures for the eastern rain forest: 

Primary forest 1 200 000 ha 

Exploited forests 3 570 000 ha 

Inaccessible areas 1 650 000 ha 

Reserves 535 000 ha 

Total 6 955 000 ha 



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Flora, vegetation and forest cover 

This, however, exceeds the total estimated by Guichon (1960) for 1958 (6 150 000 ha), without 
taking into account forest destruction since then (see below); this is likely to be because the 
FAO/UNEP category of exploited forests includes some component of Guichon's degraded 
forest. 

Battistini and Verin noted in 1972 that primary rain forest was by then found in only a 
relatively few places - the most extensive areas being in the Maroantsetra and Antongil Bay 
regions, though also on the steepest slopes of the great escarpment and in the Tsaratanana 
Massif. It is worth noting, however, that according to Le Bourdiec (1972) the southern flank 
of the Tsaratanana Massif appeared to be one of the most heavily eroded areas on the island. 

ii. Forest destruction 

The principal agent of destruction is 'tavy' (slash-and-burn) cultivation, carried out by 
subsistence farmers. Areas are cut, the vegetation is dried and fired some months later. The 
ground is then cultivated, mainly with dry land rice, but with also with maize, manioc and 
other crops, for a year or sometimes two and is then left fallow with the process repeated 
elsewhere. Degraded vegetation types (secondary forest, savoka) regrow on the land which is 
then normally recleared after an average interval of 10 years (i.e. intervals of 6, 10 or 15 
years). This cycle is repeated; however, progressive deterioration of soil structure and nutrient 
content leads to the regrowth vegetation becoming more degraded after each clearance, until 
finally the area becomes grassland or bracken-covered. Chauvet (1972) estimated that a 
maximum of 10 to 15 clearances was possible on any given site until this stage was reached, 
though the actual number will depend on the interval between clearances, the slope of the 
ground, the soil type and other factors. Tavy cultivation is often practised on very steep slopes 
(often over 20%) when the risk of erosion becomes very great. 

A gross estimate was made in the mid-1960s of 150 000 ha of forest cleared each year. Taking 
a generous estimate of 15 possible cultivation cycles on any given site, this leads to a minimum 
total loss of forest (i.e. degeneration to uncultivable land) of 10 000 ha per annum. This rate 
has almost certainly increased over the past fifteen years, owing to both the rapidly growing 
population and the ever-decreasing area of easily accessible cultivable land, which leads to 
shorter intervals between clearances on individual sites. It has also been pointed out that 
farmers always prefer to clear virgin or near-virgin land when the choice arises as this will 
ensure as rich a soil as possible. FAO/UNEP (1981) gave a figure of 40 000 ha of previously 
undisturbed closed forest cleared per year for the years 1976-80, and projected 35 000 ha for 
the years 1981-85; the great majority of this is expected to be in the eastern forests. Guichon 
(1960) estimated an area of perhaps 3 600 000 ha of savoka in 1958 although the 1981 
FAO/UNEP report quotes 2 235 000 ha, despite the fact that its area must have increased 
considerably in the intervening period. 

Clearance is greatest around settlements and in accessible areas, though with increasing 
population pressure, more and more regions are cleared. The great majority of the eastern 
coastal plain, the most densely populated part of the island (along with part of the central 
highlands), has been either cleared for permanent crops such as coffee or has degraded to 
savoka which is often very poor, consisting only of bamboo or Ravenala. 



Western Forests 

i. Area 

Guichon (1960) estimated 2 600 000 ha surviving in 1958, with around 590 000 ha of this on 
calcareous soils (mainly limestones). 



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An environmental profile of Madagascar 

FAO/UNEP (1981) quotes: 

'Virgin forest' 500 000 ha 

'Exploited forest' 1 500 000 ha 

'Degraded massifs' 650 000 ha 

Reserves 495 000 ha 

Total 3 145 000 ha 

The most recent assessments (early 1970s) indicate a few large areas of western forest 
remaining, it otherwise persisting in small isolated residual patches; overall it survives mainly 
in areas inaccessible to tribesmen or unsuitable for clearing, usually because of edaphic factors. 

It was noted in 1972 that dense forests occupied relatively large areas on limestone slopes and 
also on the sandy soils along the coast below a height of 300 m. Battistini and Verin (1972) 
reported extensive forest surviving on the Antsingy of Maintirano, which covers the dip slopes 
of the Bemaraha limestone cuesta (part of this the area is in the Reserve Naturelle Integrale de 
Bemaraha), while Koechlin (1972) additionally noted the limestone plateaux of Ankarana, 
Kekifely and Mahafaly as forested (though this last is normally included as part of the 
Southern Domain). The Ankarafantsika forest clothed the dip slope of the cuesta in the Upper 
Cretaceous sandstones and basalts of the Mahajanga region, though it was reported as only 
intact in the north (in the Reserve Naturelle Integrale de l'Ankarafantsika). Further south, the 
forest of Zombitsy near Sakaraha survived on hard-pan red sands which were unsuitable for 
shifting cultivation; in 1962 this forest was recorded by the Service des Eaux et Forets as 
21 500 ha in extent. Battistini and Verin (1972) noted that it showed the features of a remnant 
of previously much more widespread forest cover and was undergoing gradual attrition as 
outlying trees suffered from the annual grass fires on surrounding areas - this is likely to be a 
virtually universal phenomenon in the surviving western forests (see below). FAO/UNEP 
(1981) also notes the following areas as forested: Bara, Manasamody, the Bongolava Massifs, the 
area between Analalava and Ambato-Boeni and between Antsalova and Mangoky, as well as 
along the Mangoky river. 

ii. Forest destruction 

The principal threat to the western forests is undoubtedly fire, much of it deliberately started. 
The western region is the principal stock-raising area of Madagascar and fires are set each year 
to encourage new grass growth for grazing, normally during the dry season when the vegetation 
is particularly vulnerable. It has been stated that a third of the land area of the island is burnt 
every year, and in 1969 an estimated 2 000 000 ha of woody vegetation were burnt, though 
only 2000 ha of this were said to be strictly forest, the rest being shrubby vegetation or bush 
(these figures apparently refer to the west). 

The western forest is far more susceptible to fire than the east, rapidly giving way to bush and 
then wooded savannah, with a few fire-resistant palms and trees, and eventually forming 
sterile grassland, a formation which already covers perhaps 75% of the island. Annual burnings 
and grazing by livestock (principally zebu cattle, of which there are some 10 000 000 on the 
island) prevent significant regeneration taking place and it seems certain that in many areas 
degradation of soil has proceeded to such an extent that no natural regeneration is possible 
even without further disturbance. FAO/UNEP (1981) estimated net degradation of western 
forest to have been perhaps 200 000 ha since 1955. As noted above, the great majority of the 
western forest has already been destroyed and that which survives tends to be in areas more 
resistant to fire; however, as land is progressively overgrazed and population pressure increases 
even these areas come under threat, undergoing wholesale burning or suffering annual attrition 
as trees at the edge are burnt and the savannah extends - viz. the figure given above of 
2000 ha of 'true forest' burnt per annum. 

In addition FAO/UNEP (1981) estimated a net degradation through fire of 1000 ha of 
productive savanna and 4000 ha of unproductive savanna per annum, supposing that a certain 



-34- 



Flora, vegetation and forest cover 

amount of natural regeneration (to bush) took place. They thought that perhaps 50 000 ha was 
recolonized annually by shrubs, almost all of this in the west. 

Although fires for livestock grazing are undoubtedly the most important factor in the 
destruction of these forests, much land has also been cleared for crops such as maize or 
groundnuts, usually for shifting cultivation. FAO/UNEP (1981) notes that from 1936 to 1948 
large areas were cleared for the cultivation of maize as a result of policies of the colonial 
administration; ecological consequences were disastrous and policies were reversed in the 1950s, 
with a resultant marked decrease in the rate of clearance for maize. 

Land so cleared is usually rapidly exhausted and is often replaced directly by bush or 
grassland, unlike in the east where several cycles of secondary forest formation are possible; 
reafforestation then becomes a very lengthy or non-existent process. 



Southern Forests 

Guichon (1960) estimated some 2 900 000 ha in 1958, which is similar to that quoted in 
FAO/UNEP (1981) (2 990 000 ha, also referring to the 1950s). 

This is generally agreed to be the least deteriorated of the island's climax formations, especially 
on the extensive limestone lithosols which are unsuitable for agriculture, such as the plateau 
covering a large part of the Mahafaly territory and the area to the north of the Onilahy river. 
Human population is generally low in the region (though it is increasing, as everywhere on the 
island) and the vegetation is relatively unaffected by fire, combustible material being scarce. 

However, large areas have been cleared, for example in the region south of the Androy where, 
according to Battistini and Verin (1972), the Alluaudia forest existed only in patches in the 
region of Tsihombe and on the lower Mandrare. Clearing is carried out for production of food 
or cash crops such as castor-oil and sisal; much of it, such as that practised by the Mahafaly 
and Masikoro peoples, is in the form of shifting cultivation with large areas 

burnt and drought-resistant sorghum or maize sown in the ashes. Cleared areas usually form 
grassland which is quickly grazed bare. Continuing grazing pressure on many disturbed areas 
prevents, or at least hampers, regeneration and Koechlin (1972) has noted that the floristic 
composition of areas that do recover tends to differ from that in undisturbed forests. In the 
Androy and Mahafaly districts, however, the main staples of subsistence - cassava, maize, 
beans, and sweet potato - may be grown around the villages in permanent fields enclosed by 
hedges (Nelson et al., 1973). The collection of wood for conversion to charcoal for fuel is 
considered perhaps the principal threat to the southern forests and appears to be increasing 
rapidly in extent, although virtually no quantitative data are available (Richard et al., 1985). 
Charcoal is both used locally and taken for sale to major population centres. Charcoal 
production is theoretically controlled by a licensing system, with a small annual tax payable, 
but the extent to which this is enforced is unclear (O'Connor pers. comm., 1986). 'Fantislotra' 
(Alluaudia) is also collected for use as building material and Adansonia trees are felled to 
provide moisture for livestock from their swollen trunks. 

There do not, however, appear to be any overall estimates for the rate of destruction of the 
natural vegetation of the south. 



Central Highlands 

The Hauts Plateaux region in the centre of Madagascar is the area which has suffered the 
greatest deforestation. Here the forest has been reduced to a few scattered remnants, such as 
that on the Ankaratra Massif south of Antananarivo and the Ambohitantely forest on the 
Ankazobe tampoketsa north of the capital (though the former should be more correctly 
considered a montane forest, discussed below). The latter was estimated in 1964 as covering 



-35- 



An environmental profile of Madagascar 

perhaps 3000 ha, of which 2000 ha was in one continuous forest, the rest being in small, 
scattered tracts at the heads of the valleys. These areas were all recorded by Bastian (1964) as 
rapidly disappearing, mostly through the annual grass fires eroding the forest edges. This area 
was reported in 1969 as being developed as a Special Reserve of around 15 130 ha in extent. 



Montane Vegetation 

FAO/UNEP (1981) considered that the area of montane shrub (above 2000 m altitude) had 
remained roughly constant at around 600 000 ha. This vegetation is present on the large 
isolated massifs on the island - Tsaratanana and Marojejy in the north, Ankaratra in the centre 
and Andringitra in the south-east. While most of these areas are unlikely to be used by local 
people, they are extremely susceptible to fire and it seems likely that some accidental or natural 
burning will have taken place in the last 30 years. 



III.4. FOREST EXPLOITATION 

Timber 

Madagascan forest is relatively poor from an economic point of view. The area of 
commercially usable timber trees is small, and even here many individual trees have rotten or 
hollow trunks. Large areas are virtually unexploitable because of the steep terrain. Elsewhere 
transportation of timber is extremely difficult - there are no open public roads facilitating 
forest exploitation. 

Other than in cases of established land-use rights, exploitation is authorised by permit which is 
allotted either by adjudication or by private contract for a fixed time through payment of dues 
based on the land area or on the amount of production. Permits are allocated to public services 
or individuals in regions where there are no existing foresters capable of satisfying local needs. 
Produce resulting from these permits is not allowed to be commercially exploited. The 
Direction des Eaux et Forets itself apparently only exploits some reforested areas. Although 
permits are apparently accompanied by sylviculture plans, control of exploitation remains 
slight. Forest exploitation lots are scattered along or near communication routes and vary in 
size from a few dozen to some thousands of hectares. Trees currently exploited include species 
of Dalbergia, Canarium, Cryptocarya, Ocotea, Sideroxylon, Hemandia, Vernonia, Symphonia, 
Afzelia and Calophyllum. 

The Service des Eaux et Forets records 402 000 m 3 of rough timber exploited in 1979 
(probably based on the number of permits in force in that year). From the same source, the 
return per ha was around 30 m 3 for eastern forests and 10 m 3 for western forests, giving 
9100 ha of exploited eastern forest, 4200 ha of western forest and 700 ha of southern forest. 

Most exploitation uses traditional methods: felling is by axe, and wood is extracted and cut into 
planks or cross-sections with a long-saw or axe. Yield is very low (ca 20%). There is little 
mechanised exploitation. Here, felling is carried out either with axes or chain-saws, and 
trimming with chain-saws. Timber is extracted, as rough logs, which are cut up either on site 
or at the saw-mill. Use of chain-saws is not general but is spreading. The use of tractors for 
clearing and wood-peeling as much as for creation of tracks, is now fairly widespread on 
mechanised sites. Most of the relatively accessible forest has already had the best timber 
systematically removed and there are virtually no remaining areas where the standing crop is 
economically viable. 

Since 1975 no valuable unworked wood (e.g. of Dalbergia) can be exported; only worked 
timber (de-barked, cut into planks) is allowed to leave. 

Most of the forestry industries (especially timber-mills) are in the private sector; timber-mills, 
numbering 60 at the end of 1975, are scattered throughout the island with a heavy 



-36- 



Flora, vegetation and forest cover 

concentration in the province of Antananarivo. Their capacity varies from 500 m^ per year to 
3000 m , with 80% having a capacity lower than 1500 m^ per year. 

Other forest products 

The rural population exercises rights of usage which allow them to freely obtain wood for the 
construction and preparation of dwellings, firewood and other products for their own use. 
Theoretically, these should be under the control of permits issued without charge though in 
practice exploitation is often carried out without them. FAO (1978) estimated 400 000 np of 
'bois de service 1 (timber for building), 5 200 000 m^ of firewood and charcoal (wood 
equivalent). 

A 1980 Direction des Eaux et Forets report gave annual production of 2 520 000 tonnes of 
fire- wood and 71 000 tonnes of charcoal. 

Honey, beeswax, resins and raphia (4225 tonnes according to the above) are important miner 
forest products. 



III.5. REAFFORESTATION 

Reafforestation projects have been undertaken for several decades, especially in the Hauts 
Plateaux. One of the earliest on any scale apparently was the establishment of broadleaves, 
especially eucalyptus, along the railway from Antananarivo in 1910 to provide fuel- wood for 
the locomotives (since the introduction of diesels, these plantations have been used for other 
purposes). Most plantations have been of eucalyptus for firewood and local building, though 
conifers have been increasingly used since the 1950s following success with Pinus patula. 

FAO/UNEP (1981) distinguishes between industrial and non-industrial plantations: 

i. Industrial plantations - estimated 112 000 ha 

These are very largely based on pines, some for the production of industrial timbers, others for 
pulp-wood. Species used include Pinus patula, P. kesiya, P. pinaster and Cupressus lusitanica. 
There are some plantations of mixed pines and cypress and of pines and eucalyptus, and also 
very limited areas of other timber trees such as Afzelia bijuga, Terminalia superba, Cedrela 
odorata, Tectonia grandis, Eucalyptus spp. and other species. 

In 1980 it was envisaged that 35 000 ha of industrial plantations would be planted between 
1981 and 1985, of which 85% would be conifers and 15% broadleaf trees. 

ii. Other plantations - estimated 154 000 ha 

These are principally for firewood and 'bois de service'. Different Eucalyptus species are used, 
including E. grandis, E. camaldulensis, E. saligna. E robusta, E. viminalis, E. racemosa, E. 
citriodora, E eugenioides and the hybrid E. 12 ABL, either in monocultures or mixed 
with Acacia mearnsii, A. dealbata, A. melanoxylon, Casuarina equiseti folia, Terminalia superba, 
Afzelia spp, Tectonia grandis or conifers. 

In 1980 it was envisaged that 25 000 ha of non-industrial plantations would be planted between 
1981 and 1985, of which 20% would be conifers and 80% broadleaf trees. 

According to the 1981 FAO report about three quarters of plantations are state owned; 15% 
belong to local communes and 10% are private. These figures are based on a questionnaire sent 
out in 1973 and can be expected to be approximate at best. 

Important reafforestation projects undertaken since 1968 by the Direction des Eaux et Forets et 
de la Conservation des Sols include: 



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An environmental profile of Madagascar 

- Industrial plantations in the Haute Matsiatra, covering in 1976 an area of around 35 000 ha, 
almost totally planted with Pinus patula; 

- Industrial plantations in the Haut Mangoro, covering in 1976 nearly 50 000 ha of Pinus kesiya\ 

- Manankaso, where plantations of Pinus patula totalling some 10 000 ha were planned; several 
hundred hectares had been planted by the end of 1976; 

- Plantations in the Vakinankaratra, a region which could be joined to that of the Haut 
Mangoro to supply the same industry; in 1969, 1200 ha of Pinus patula were planted. 

- Planting of cashew Anacardium occidentale in Mahajanga and Antseranana provinces. The 
objective, apparently attained by 1981, was the planting of 40 000 ha of cashews to supply 
factories processing the nuts. By 1969, 11 000 ha had been planted around Mahajanga, 
10 000 ha at Ambilobe, and around 9000 ha scattered through the two provinces. 



- A scheme in the Morarana area which envisaged the planting of softwoods to supply a match 
factory at Moramanga as well as other industries. The final aim was the creation of 1500 ha of 
poplars, of which 350 ha had been planted in 1969 (in 1984 it was reported that the factory 
had closed (Jolly el al., 1984)). 

- A scheme to provide wind-breaks in the south of the island which aimed to plant some 
100 km per year. 

No significant replanting using native species has been undertaken, though small scale 
experimental planting has been undertaken at some forest stations. 



III.6. ETHNOBOTANY 

Introduction 

Madagascar is currently a net importer of pharmaceuticals (World Bank figures of 3664.5 
million Madagascan Francs for 1977, equivalent to ca U.S.S14 500 000). In 1977, the World 
Health Assembly of the World Health Organization (WHO) adopted a resolution urging 
governments to promote interest and research on their traditional medical systems. The 
rationale behind this was that, where proven effective, traditional medicine could provide an 
important health care resource in developing countries since it is both culturally acceptable and 
economically feasible (Akerele, 1984). Furthermore, medicinal plants of proven effectiveness 
could theoretically be cultivated and exported, thereby reducing import expenditures while 
increasing employment opportunities and generating foreign exchange (Plotkin, 1982). WHO 
has established Collaborating Centres for Traditional Medicine in Ghana, Mali, Nigeria, Sudan, 
India, China, Italy, Mexico, and the United States (Gyllenhaal, 1985). Appreciating the 
richness and uniqueness of its cultural heritage as well as its flora, the Malagasy government 
has expressed a strong interest in research on traditional medicine and medicinal plants, and 
has established a Department of Ethnobotany within the Centre National de Recherche 
Pharmaceutique (C.N.R.P.). 



The potential value of the Madagascan flora 

The Madagascan Rosy Periwinkle {Catharanthus roseus) belongs to the family Apocynaceae, 
whose species are often rich in alkaloids. C. roseus is the source of over 75 alkaloids, two of 
which are used to treat childhood leukemia and other cancers with a high rate of success. 
Other experimental pharmacological activities - diuretic, hypoglycaemic and antiviral - have 
also been associated with alkaloids extracted from this species (Svoboda and Blake, 1975). It 



-38- 



Flora, vegetation and forest cover 

should be noted that C. roseus was first subjected to laboratory investigation because of its use 
by local peoples as an oral hypoglycaemic agent (Cordell and Farnsworth, 1976) 2 . Sales of 
these anti-cancer drugs worldwide in 1980 were estimated to be worth approximately 
$50 million wholesale prior to 100% retail mark-up (International Marketing Statistics, 1980, 
quoted in Myers, 1984; Svoboda, 1981; Myers, 1984). Myers (1984) forecasts that demand for 
these drugs will increase by 15% by a year. It is indeed significant that Madagascar received 
nothing in royalties from these sales, since the drugs are currently extracted from Rosy 
Periwinkles cultivated in other countries (Myers, 1984). Furthermore, there are several other 
species of Catharanthus on the island and one of these, C. coriaceous, is considered to be in 
"utmost danger" of extinction (Thompson, 1984). 

The actual and potential utility of the Madagascan flora is not limited to the field of medicine. 
Two of the important agricultural commodities on the island are coffee {Coffea arabica) and 
vanilla {Vanilla plant folia). In 1980, Madagascar produced 80 000 tons of coffee, which 
amounted to 35% of the dollar value of the country's entire exports for 1981 (see Part II). 
Coffee exports in 1982 were worth 34.91 billion Madagascan francs (International Monetary 
Fund, 1985). Although C. arabica originated in Ethiopia, Madagascar is home to a number of 
wild species, perhaps as many as fifty (Guillaumet and Mangenot, 1975). Some of these species 
may have potential in commercial breeding since they produce beans with little or no caffeine 
(Guillaumet, 1984). As far as we could ascertain, none of the Madagascan Coffea germplasm 
has been collected to conserve them for future genetic work. 

Next to petroleum, coffee is one of the most important commodities in international trade and 
is a mainstay of the economies of several tropical countries (Imle et al., 1977). Nonetheless, 
commercial coffee (primarily Coffea arabica) is rather susceptible to certain fungal diseases. 
The spores of these fungi are very short lived, yet rapid modes of transportation have already 
resulted in at least one introduction of the disease from the Old World to the New World 
tropics (R.E. Schultes, pers. comm.). As a result, both Central American and Brazilian coffee 
crops were attacked by a rust disease in the 1970s. Fortunately, a rust-resistant strain from 
Ethiopia was introduced with successful results. Not only is the continued success of 
commercial coffee cultivation dependent on the conservation of wild relatives, but coffee 
germplasm must also be maintained in living collections, since prolonged seed storage is not 
feasible (Ferwerda, 1976). Guillaumet (1984) reported finding five different coffee species 
within a 2 km area in a lowland rain forest reserve, and he claimed that most Coffea species in 
Madagascar exist only in reserves and protected areas. 

Artificial pollination of commercial vanilla (Vanilla planifolia was first developed in the late 
1800s, making cultivation possible in many areas, including Madagascar (Correll, 1953). 
Vanilla plantations were first established on Nosy Borah (He Ste Marie) and Nosy Be, and were 
then initiated on the mainland (Koechlin et al., 1979). Today, Madagascar produces most of 
the world's commercial vanilla: in 1982, vanilla exports were worth 16.76 billion Madagascan 
francs (International Financial Statistics, 1985). Despite the fact that cultivated vanilla has a 
very narrow genetic base (G. Wilkes, pers. comm.) and that commercial vanilla plantations on 
the island are periodically ravaged by fungal diseases (Guillaumet, 1984), we were unable to 
learn of any attempts to collect, preserve, or otherwise employ the germplasm of the 5-6 
species of wild Vanilla endemic to Madagascar. 

Madagascan fibre plants like the Raphia palm should also be investigated for their commercial 
potential. Rattan from South-east Asia is the basis of a multimillion dollar industry, yet 
overcollecting has severely reduced available stands in some areas (Dransfield, 1981; Myers, 
1984). 



2 Interestingly, its use was originally reported from Jamaica; nevertheless, records of its use as 
a folk medicine in Madagascar exist as well (see Appendix 5). 



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An environmental profile of Madagascar 

Previous studies of ethnobotany in Madagascar 

More than seventy books and papers dealing with at least some aspect of the useful flora of 
Madagascar have been published to date. As far as we are able to ascertain, only one of these 
publications (Debray et al., 1971) was based on field study which included collection of 
voucher specimens. The great majority of the works are either taxonomic or anthropological 
and demonstrate a paucity of first-hand ethnobotanical information. 

This statement does not intimate, however, that there are not many useful data available. 
Excellent compilations on the ethnobotany of Madagascar (e.g. Heckel, 1910; Pernet and 
Meyer, 1957; Boiteau, 1974; etc.) contain information on the utility of hundreds of local 
species. Vernacular and scientific names as well as actual uses are presented, but there are no 
voucher specimens, no methodologies and sometimes no method of preparation given. The 
most useful works on the ethnobotany of Madagascar include (in chronological order): 

1) Heckel, 1910 - Heckel presents a compendium of ethnobotanical data from various 
peoples of Madagascar. Although most of the information seems to be based on the use of 
plants by the Merina, utilization of species by other tribes (including the Betsileo, 
Betsimisaraka, and the Sakalava) are also presented. This book contains data on names 
and/or uses of over 700 species of plants, and entries are cross-referenced by both 
scientific and vernacular names. 

2) Dandouau, 1922 - This paper on the charms and remedies of the Sakalava and Tsimihety 
notes that these tribes believe that all sickness is caused by spirits. Treatments are 
specified for all of the maladies listed, although unfortunately the plants are for the most 
part referred to only by vernacular name. The author notes, however, that Aphloia 
theaeformis and Cassia occidentalis were being prescribed by medical doctors at that time 
to treat both common and hepatic fevers. 

3) Linton, 1933 - Linton's monograph on the Tanala contains a section on alcohol and 
narcotics employed by members of the tribe. 

4) Dubois, 1938 - This monograph of the Betsileo contains an important contribution to 
Madagascan ethnobotany. The study features a section on the plant medicines of the 
tribe, including alphabetical listings of both the illnesses and the plants employed in their 
treatment. Fifty of the plants discussed are identified by their scientific name. 

5) Terrac, 1947 - Terrac's thesis represents the second major compilation of the medicinal 
plants of Madagascar. The work includes data on some 800 phanerogamic and 30 
cryptogramic species. 

6) Robb, 1957 - A little known paper which presents an excellent overview of the use of 
ordeal poisons in both Africa and Madagascar. The author lists three species employed in 
Madagascar: 

a) "Tanghin" - Tanghinia venenifera (Apocynaceae). "Tanghin" was employed as an 
ordeal poison in the judgement of all crimes, especially sorcery. The nut or kernel of 
this tree of the eastern forests is highly toxic, containing a cardiac glycoside, the 
physiological properties of which are similar to those of strophanthin and ouabain, the 
active principles of an ordeal poison employed in West Africa. The use of "tanghin" to 
find and kill sorcerers once resulted in the death of 6000 people. It is still employed in 
some areas for the purposes of assassination (D. Gade, pers. comm.). 

b) "Ksopo" - Menebea venenata (Asclepiadaceae). "Ksopo" was the ordeal poison of the 
Sakalava and is found in their arid tribal homeland in the west/north-western part of 
the island. Like the aforementioned "Tanghin", the toxicity of "Ksopo" is attributed to 
the presence of a powerful cardiac glycoside. 



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Flora, vegetation and forest cover 

c) "Couminga" - Erythrophlewn couminga (Leguminosae). The toxic bark of "Couminga" 
was employed as an ordeal poison throughout Madagascar and the Seychelles. So 
poisonous did the locals consider this species, that they claimed that the fragrance of 
its flowers, the rain water that washed the leaves, and the smoke from burning parts of 
the plant all would be fatal; this is reminiscent of the stories told of the "upas" tree 
(Antiaris toxicaria) of tropical Asia (R.E. Schultes, pers. comm.). As in the two 
preceding examples, the active principles of "couminga" were cardiac glycosides. 

Although the article concludes that the use of ordeal poisons was not practised after 1920, 
current reports suggest that these rituals may still persist in some of the more remote areas 
of the island. 

7) Pernet (1957, 1959) - These works are regarded as the forerunner of the pharmacopeia of 
Madagascar. They draw on the research conducted at the Laboratoire de Chimie Vegetale 
in Madagascar between 1954 and 1959. 

8) Pernet and Meyer, 1957 - The Pharmacopee de Madagascar presents an excellent 
compilation of ethnomedical uses of plants. Drawing on 58 references, this work contains 
data on 1171 uses of over 450 species. Although no information is provided on vernacular 
names, use by particular tribe or chemical composition, data are cross-referenced by 
taxonomy and ethnomedicinal use. 

9) Decary, 1964 - An overview of the useful palms of Madagascar, this paper contains 
information on 31 local species and on two introduced palms. Data are presented on local 
distribution, scientific and vernacular names, fibre, fruit, edible starch, vegetable salt, 
palm hearts, and palm wine. Specific mention is made of palms used by the Sakalava and 
Betsimisaraka. 

10) Boiteau et al., 1968a, b - These papers furnish data on bitter barks used for ethnomedical 
purposes by various peoples in Madagascar. Information is offered on practical and 
economic uses, etymology of vernacular names, and particular species employed. 

11) Ratslmamanga et al., 1969 - This work represents the initial effort to publish an official 
Madagascan Pharmacopeia along the lines of the Codex Francais -- the Pharmacopeia of 
France. Unfortunately, it covers only the first 39 elements (i.e., it does not go beyond the 
letter "A"), and no subsequent portions appear to have been published. 

12) DeBray et al., 1971 - Published by ORSTOM, this book represents the single most 
important primary reference yet published on the ethnomedicine of Madagascar. The first 
section deals with the plants of the south-west and their use by the Bara, Mahafaly and 
Sakalava. Section two offers data on the plants of the east coast -- those employed by the 
Antaimoro and the Betsimisaraka. The last section is the largest of the three and details 
the plants of the plateau employed by the Merina. 

The importance of this work is multifold: it is all based on original fieldwork; it features 
exact collecting localities and voucher specimens; it offers vernacular and scientific names; 
and it gives the results of chemical analyses of the useful species. On the negative side, it 
makes virtually no reference to earlier works on the ethnobotany of the peoples or the 
plants. Furthermore, none of the tribes studied (with the possible exception of the 
Merina) was examined in depth. Nevertheless, this work represents a benchmark for 
future field and laboratory research on the ethnobotany of Madagascar. 

13) Rarafindrambao, 1973 - A study of Buxus madagascarica, a plant which is employed for 
a variety of purposes in Madagascar. 

14) Taylor and Farnsworth, 1975 - Although much has been written on the 
genus Catharanthus, this book offers one of the best overviews available, featuring data on 
the botany, distribution, taxonomy, chemistry, phytochemistry, pharmacology, and 
synthesis of Catharanthus alkaloids. Chapters include: 

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An environmental profile of Madagascar 

1. A Synopsis of the Genus Catharanthus (Apocynaceae) by William T. Steam 

2. The Phytochemistry and Pharmacology of Catharanthus roseus (L.) G. Don. by 
G. Svoboda and D.A. Blake 

3. The Phytochemistry of Minor Catharanthus species by M. Tin-Wa and N.R. Farnsworth 

4. Structure Elucidation and Chemistry of the Bis Catharanthus Alkaloids by Ronald 
J. Parry 

6. Tissue Culture Studies of Catharanthus roseus by David P. Carew 

7. Biochemistry of Dimeric Catharanthus Alkaloids by William A. Creasey 

8. Clinical Aspects of the Dimeric Catharanthus Alkaloids by R.C. DeContin and 
W.A. Creasey 

15) Boiteau, 1974-79 - Boiteau was a French agronomist who founded the botanical garden 
and zoo at Tsimbasasa, currently under the Directorship of one of the co-authors of this 
report. In 1974, Boiteau began publishing the Dictionnaire des Noms Malgaches des 
Vegetaux in serial parts in the Italian journal Fitoterapia. The entries were filed 
alphabetically by vernacular name, and it was originally envisioned that this work would 
include virtually all of the available information on the useful species of the island. 
Unfortunately, Boiteau died in 1980 and the Dictionnaire reached only the letter "L." 
Boiteau's wife and daughter assembled the data posthumously, and the second half of the 
work still exists in typescript. Since the journal Fitoterapia is of limited distribution and 
as the Dictionnaire is of rather limited use in its current half-published form, it is strongly 
suggested that the entire manuscript be published as a complete .vork with financing from 
an international conservation organization or botanical garden. 

16) Boiteau and Potier, 1976 - A general overview of the importance of ethnomedical 
knowledge, this article cites examples from Madagascar (such as the 
anti-cancer Catharanthus roseus and the biodynamically active species of Evodia employed 
in circumcision rites). 

17) Ratsimamanga, 1977 - A general overview by the noted Madagascan ethnobotanist 
Rakoto Ratsimamanga of the potential of ethnomedical species for the production of local 
pharmaceuticals. 

18) Boiteau, 1979 - Information on Madagascan plants specifically organized for doctors 
wishing to use local resources for medicinal purposes. This work contains a breakdown of 
useful plants by category of use (insecticide, purgatives, febrifuge, etc.) with over 90 
categories, a list of simple tisanes and their uses, and other recommended preparations and 
their uses. 

19) Scarpa, 1980 - This paper is yet another overview of the importance of traditional 
medicine, offering a number of examples from Madagascar. The author concludes that 
traditional treatments should not be discarded but rather blended with modern scientific 
knowledge, hopefully with holistic results. 

20) Rabesa, 1985 (unpublished) - Dr Zatera Antoine Rabesa, Minister of Scientific Research, 
recently completed an ethnobotanical study of the people living near Lac Alaotra 
(Sihanaka?); it has not yet been published. Dr Rabesa is currently in the process of 
initiating a similar study of the Sakalava living near Mahajara. It is this type of research, 
conducted by local scientists, that bodes well for the future of ethnobotany in Madagascar, 
and that should be financially and logistically supported by international conservation 
organisations, pharmaceutical companies, and botanical gardens and museums. 



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Flora, vegetation and forest cover 

Appendix 5 contains a preliminary ethnobotany database for Madagascar, detailing plants and 
their medicinal uses, classified both by plant species and by use. 



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PART IV. MARINE AND COASTAL ECOSYSTEMS 

IV.l. RELIEF 

Littoral and coastal relief is characterized by marked east-west asymmetries in the same 
manner as the continental relief (Battistini, 1972; Guilcher and Battistini, 1967); these have 
strong influence on the distribution of coral reefs and mangroves, Madagascar possessing 
significant areas of both. 

The continental shelf is very narrow on the east coast, with the 100 m depth contour 3 to 5 km 
offshore and the 1000 m contour 15 to 30 km offshore; by contrast, on the west coast the 
100 m contour is 30 to 80 km offshore, the 1000 m contour 40 to 150 km. The shelf is notably 
wide between Cap d'Ambre and Mahajanga, between Soalala and Morondava, and between 
Taolanaro and Androka. 

Tidal amplitude differs considerably - on the east coast high-tide level is very low (some 
50 cm), while on the west it commonly reaches 3 to 5 m. This (and the very wide extent of the 
western coastal plain) allows the development of extensive brackish water areas in the west, 
while in the east the transition from sea to fresh water is usually very rapid. 

Following Battistini (1972), littoral relief can be divided into four main regions: 

i. The north-west between Cap d'Ambre and Cap Saint-Andre shows a characteristic 
submerged continental relief and has the most indented coastline of the island, with many bays 
and capes (often of relatively high elevation) and large numbers of offshore islands. 

ii. Along the west and south-west between Cap Saint-Andre and Cap Sainte-Marie, the 
coastline is low-lying and monotonous in appearance, being more or less without indentations 
and having very long sandy beaches, with mangrove swamps behind. There are many large 
deltas, only the largest of which (e.g. Mangoky and Tsiribihina) extend for a short distance out 
to sea; others are aborted to a greater or lesser degree by ocean swell shaping a straight 
coastline. This contributes to the formation of a large, low-lying littoral plain. 

iii. The extreme south is characterised by considerable quaternary dune accumulations; 
the oldest of these are sandstone with cross bedding, and lead to a rocky coast in many 
regions. Elsewhere (such as along the Mahafaly coastline in the south-west) there are vast 
sandy beaches. No mangrove swamps of any size are found. 

iv. The east coast between Taolanoro and Foulpointe is more or less straight for over 
700 km and is low lying with a continuous offshore sandbar, behind which is a string of 
lagoons linked together by the artificial Canal des Pangalanes. North of Foulpointe relief is 
more varied, and becomes quite high in the Mananara region, in Antongil Bay and around Cap 
Masoala. 



IV.2. MANGROVES 

Madagascar possesses the largest areas of mangrove in the western Indian Ocean. In 1972 
Keiner estimated there to be ca 330 000 ha of mangroves on the island; at least 321 000 ha of 
this was along the western coast. On the east there were only eleven mangrove sites of any 
size, the largest of these being ca 2220 ha in Rodo bay. 

On the west coast mangroves could be divided into some 29 separate areas. Seven of these 
were over 20 000 ha in extent, with the largest, in the Bay of Bombetoka, covering some 
46 000 ha. In total, there were 320 000 ha of mangroves in areas of over 1000 ha with an 
additional 10 000 ha of small or very small areas of mangrove often at the mouths of streams 
or in small bays. Of particular note are the 18 000 ha of mangrove in the Loza, a very large 
lagoon in the north-west of the island. 



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An environmental profile of Madagascar 

Floristic composition and zonation are typical of Indian Ocean mangrove systems. The 
following widespread species reportedly occur: Rhizophora mucronata, Avicennia marina, 
Sonneratia alba, Ceriops tagal, Bruguiera gymnorrhiza, Xylocarpus granatum, X. moluccensis, 
Lumnhzera racemosa and Heritiera littoralis. The Ceriops is sometimes considered an endemic 
species, C. boiviniana. 



Economic importance 

Mangroves are very important for inshore fisheries, serving as nursery areas for many species 
of fish and Crustacea. There are otherwise few data on utilization of mangroves, though it is 
noted that generally the most extensive mangrove areas are in regions of low population density 
which are inaccessible for part of the year (Keiner, 1963, 1965, 1972). 

Amongst Crustacea, Penaeid shrimps (especially the large species Penaeus monodon and P. 
indicus) and the crab Scylla serata are important (see Part V.10), and among fish the families 
Carangidae, Carcharinidae, Mugilidae, Serranidae, and Sparidae. 

Fisheries of sharks and saw-fish are important in all mangrove areas; salted-dried meat is the 
object of many transactions, especially in the region of Mahajanga. 



IV.3. CORAL REEFS 

Reefs on Madagascar are extensive and numerous and include good examples of almost all of 
the main classical reef types. Some of these have been very well studied; the barrier reef at 
Toliara (see separate account) has probably been studied more than any other reef in the Indian 
Ocean, while others, particularly on the east coast, remain largely undescribed. Most of the 
work done on Madgascan reefs has been from the French group led by Battistini, Pichon and 
Picard, and much of the following is derived from the extensive bibliography of this group (see 
'References'). Most information is taken from Battistini (1960 and 1964), Clausade et al. 
(1971), Pichon (1971a, 1972a, 1972b, 1974 and 1978b) and UNEP (1982b). Bibliographies of 
marine studies up to the early 1970s are given in ORSTOM (1973 and 1975). Most research has 
been carried out at Nosy Be in the north-west and Toliara in the south-west where the French 
authorities maintained research stations. Research still continues in these areas, the stations 
having been handed over to the Madagascan authorities. 

Regional variations in water conditions influence the distribution of reefs. Tidal amplitude 
differs considerably. On the east coast high-tide level is very low (some 50 cm), while on the 
west it commonly exceeds 3 m at spring-tide. Western tides are regular semi-diurnal, with low 
tides at noon and midnight. Currents also vary regionally. They are predominantly southerly 
on the east coast, where they are derived directly from the South Equatorial Current; on the 
west coast they are usually northerly, derived from either the South Equatorial Current as it 
flows around the south of the country or from a counterflow to the Mozambique Current. On 
reefs which have been well studied, local current patterns have been found to be complex 
because of the large and irregular expanse of many of the reef areas and the large tidal range. 
Surprisingly, the temperature differences between north and south (between which there is a 
difference in latitude of 10°) do not appear to have any influence on the richness of the coral 
fauna. At Toliara (Tulear) stony corals disappear at a depth of about 30 m; off Nosy Be they 
extend to depths of 45-60 m. At Toliara temperatures in the lagoon average 21.5°C in 
July/August and 30°C in January/February. Sixty-three genera of reef-building corals have 
been found in the Nosy Be region and sixty-two at Toliara where the waters are colder. 



Distribution of reefs 

Reefs of the east coast are least known. There are rudimentary but extensive fringing reefs 
behind which run a chain of shallow lagoons. Behind the reefs and a series of coastal dunes 
the lagoons were once connected to form an inland waterway called the Pangalanes Canal. This 



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Marine and coastal ecosystems 

was once navigable for about 700 km but much of it is now colonized by herbaceous plants 
(UNEP, 1982b). In the north-eastern part around Antseranana there is an emergent reef with a 
channel up to 8 metres deep (Pichon, 1972a). South of Antseranana are two superimposed 
emergent reefs, the older elevated to 16 m and partly covered by dunes. Nearer the coast this 
system becomes fragmented, and has the appearance of a chain of coral islets connected to the 
land. This series of structures includes the Leven Islands (Pichon 1972a) which are bordered at 
their outer edge by a discontinuous fringe of coral. Such coral reefs are numerous amongst the 
small islands of the east coast (Rabesandratana, 1984). Further south, Nosy Borah (He 
Sainte-Marie) has fringing coral growth, while in the Toamasina region there is a submerged 
and fragmented barrier reef. This also has coral growth on the seaward periphery, with a reef 
flat covered with seagrasses. The sand bar to the south which runs for 700 km is considered by 
Pichon (1972a) not to have reef formations, although a few coral communities may be found 
which are isolated and not true reefs. 

The west coast has the main reef formations covering a distance of more than 1000 km 
(Pichon, 1972a; Rabesandratana, 1985). These are located in the north-west (from Cap 
d'Ambre to Narendy Bay), and south-west, separated by a central region with relatively sparse 
reef growth between the Mangoky delta and Androka. In the far north-west, the coast is 
bordered by emergent fossil reefs up to 10 metres above present sea level. Living fringing 
reefs are well developed along the mainland coast and around offshore islands, except in the 
vicinity of deltas and their adjacent bays. Fringing reefs are well developed between Cap 
d'Ambre and Narendy Bay; narrow reefs occur between Courrier and Bejotaka Bays; small 
reefs are found off Cap Sebastien. Fringing reef is found on either side of Ambavatoby Bay 
and along the west coast of Ampasindanva Peninsula from the Kakamba estuary to the Bay of 
Rafaralahy. There is a long fringing reef between Ramanetaka Bay and Ansatramahavelona 
and on the west coast of Nosy Lava. Further south there are a few small isolated reefs, in 
particular north of the entrance to Mahajamba Bay, at the entrance to Mahajunga Bay, 
north-east of Boina Bay, on either side of Baly Bay and between the latter and Antaly Bay. No 
detailed studies have been made south of Narendy Bay. Fringing reefs are found around some 
of the small volcanic islands along this coast such as Nosy Vahila, Nosy Mananono, Tanykely, 
the Radama Islands, Nosy Saba, etc. Extensive coral formations are found in some of the bays 
along the coast where there is little sedimentation, e.g. Lotsoina Bay, Ampanasina Bay. The 
more exposed reefs, notably some around the island Nosy Be (see separate account), have 
typical well-developed structures, including boulder zones and spur and groove systems. The 
Baie d' Ambaro, also in the north-west, has been studied- in detail because of its rich shrimp 
stocks (Daniel et al., 1970; Daniel, 1972). Off shore in this area, coral formations occur on the 
Banc du Leven (Daniel el al., 1972). Octocorals from north-western Madagascar are described 
by Verseveldt (1973). 

In addition to these inshore and fringing reefs, there is a rise in the sea floor 10-60 km further 
off shore, at the edge of the continental shelf (Pichon 1972a). This rise may be a submerged 
barrier reef or a cuesta formed during an emergent phase. Much of it lies 5-15 metres deep or 
less in places, but is cut by several channels opposite large river deltas. At its northern extent 
parts are emergent, forming reefs which support the sand cays of Nosy Anambo, Nosy Fasy, 
Nosy Faty and Nosy Foty. Elsewhere the surface of the barrier is covered with vast sandy 
plains and has a coral cover of only 10% in most places; it is likely the reef is not growing, 
though no explanation is available for this. 

The central section of the west coast has no reefs or has poor fringing reef, although off shore 
there are two groups: the Pracel Shoal and reefs around the Barren Islands. The former are 
separated over 100 km, but only two parts, Chesterfield Island and Nosy Vato, are permanently 
above water and in both cases the island is in the north-east of the reef flat, which shows the 
influence of the dominant seas from the south-west (Pichon, 1972a). The Barren Islands 
support a greater number of emergent reefs and sand cays and extend over 50 km. It is 
possible that these are the southernmost extent of the offshore barrier described for the 
north-west part of Madagascar. Little work has been done on the reefs of the central sector, 
and further study is required. 



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An environmental profile of Madagascar 

Reefs of the south-west sector are well developed and the best known (Clausade et al., 1971; 
Pichon, 1972a, 1972b, 1978a, 1978b, Thomassin, 1978b; Vasseur, 1981; Weydert, 1973a, 
1973b). Fringing reef, barrier reefs and reefs with sand cays are all represented, together with 
intermediate types. Due to the narrowness of the continental shelf, the reef often changes 
from barrier to the fringing type. Winds and wave energy come from the south-west with 
enough strength to allow crustose coralline algae to dominate in shallow water. Sand cays are 
particulary numerous in the northern part of this sector, between the Bay of Assassins and the 
Mangoky delta. Two kinds of cay exist: those which rest on isolated parts of a barrier well 
offshore and are separated from it by up to 30 metres of water, and those located on a fringing 
reef situated behind the barrier. Both exhibit adaptation to the prevailing south-westerly seas, 
with a clearly marked succession of spur and groove system to seaward, an algal flat and a 
boulder tract and an inner reef flat with corals before the island. To leeward of the islands are 
seagrass patches, before a leeward slope with spurs and grooves. 

From the Mangoky delta there is almost continuous reef to the Onilahy River (Pichon 1972a). 
The first part of the Bay of Assasins has a series of small fringing reefs. South of this, the 
fringing reef becomes 2-3.5 km wide, extending for almost 80 km. It is interrupted by 
occassional passages, and contains a channel between it and the shore which is 10 metres deep 
in the north, shallowing to 1.5-5 metres in the south, and disappearing at Manombo. South of 
Manombo, the coral reefs move offshore opposite Ranobe Bay and Toliara Bay, although 
between these there is a fringing reef adjacent to the mainland. The reefs of Ranobe Bay are 
described by Clausade et al. (1971). Opposite Toliara is the Grand Recif which extends for 
18 km, and is up to 3 km wide (see separate account). From the Onilahy River, a fringing reef 
extends southwards almost unbroken for nearly 100 km although it is rather depauperate in 
comparison with more northern reefs. Its width varies greatly from 0.5 to 3.5 km in the 
northern part because of a very embayed shoreline. This reef disappears at Lanivato. A small 
fringing reef appears south of this at Itampolo and then at Androka. South of the latter, reefs 
only exist offshore around the sand cay of Nosy Manitsa and the Etoile Shoal (Pichon, 1972a). 
The latter are the highest parts of another submerged barrier reef on the edge of the 
continental shelf which runs for about 50 km. 



Conservation and utilisation of reefs 

A general description of marine and coastal resources is given in Anon. (1985). Reef areas, 
cays and lagoons are important habitats for many groups of animals which are epxloited 
locally, or in some cases for national or international markets, including marine turtles, the 
Dugong Dugong dugon, molluscs, crustaceans and fishes. Reef fish on the west coast are 
described by Fourmanoir (1963); other groups are fully discussed in the relevant sections of 
Part V and in Appendix 3. 

The impact of current human activities on the reefs is virtually unknown. A threat of possible 
importance to Madagascan reefs is sediment accumulation as a result of the massive erosion and 
concomitant soil runoff which affects much of the island (Pichon, 1984, and see Parts I and III). 

Sediment accumulation is acute in many areas, as illustrated by the loss of use of the port of 
Mahajanga due to the deposition of 100 million cubic metres of silt in 25 years. Elsewhere, in 
the north, river deltas have expanded and sediments have reportedly been deposited on beaches 
and reefs, changing lagoonal current patterns and having potentially adverse effects on local 
fisheries. However, there appear to be no concrete data on the effects of sediment on 
Madagascan reefs. 

Over-fishing is reportedly becoming a serious problem, especially on the Grand Recif (see 
separate account). Pollution on the western coast, where most reefs occur, appears to be of 
minor importance at present and dynamiting for fish, which is a common practice in many 
countries, is not a major threat (Pichon, 1984). Corals have reportedly been used in the past 
for building materials (Rabesandratana, 1984), though it is not clear if this continues. 



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Marine and coastal ecosystems 

There is at present no legislation to protect coral reefs, though the Grand Recif at Toliara has 
twice been proposed as a Marine Park (see separate account). It is recommended that surveys 
of the more accessible reef areas, particularly Nosy Be, Toliara and Nosy Borah, should be 
carried out to determine their status and the establishment of marine parks should be seriously 
considered (Anon., 1985). Randrianarijaona and Razafimbelo (1983) stress the need to take 
into consideration socio-economic factors in any such areas. Multiple use Marine Parks might 
be suitable, to ensure that the economic benefits to be gained from tourism are not overlooked. 

A series of projects concerning coastal zone management were presented at a conference on 
Conservation des Ressources Naturelles au Service du Developpement held in Antananarivo in 
November 1985. These included a study to evaluate coral reef resources, the establishment of 
the Pare National du Grand Recif de Toliara, and a number of projects relating to fisheries 
and pollution (Anon., 1985). 

Data for three reefs or groups of reefs of particular interest are provided below. This 
information is taken from the forthcoming Directory of Reefs of International Importance. 
Details, where available, are provided under the following headings: Name; Geographical 
location; Area, depth, altitude; Physical features; Reef structure and corals; Noteworthy fauna 
and flora; Scientific importance and research; Economic value and social benefits; Disturbances 
or deficiencies; Legal protection; Management; Recommendations. 



NAME Grand Recif proposed Marine National Park 

GEOGRAPHICAL LOCATION South-west Madagascar, near Toliara. 23°25'S, 43°40'E. 

AREA, DEPTH, ALTITUDE Grand Recif is over 15 km long from north to south, and up to 
3 km wide. Depths on the seaward slope exceed 50 m. 

PHYSICAL FEATURES The following is derived principally from the work of Clausade et al. 
(1971) and Pichon (1971a, 1972a and 1978b), Thomassin (1978b), Vasseur (1981) and Weydert 
(1973a, 1973b). 

The reefs of Toliara, including Grand Recif, are protected from the south-east trade wind by 
the mainland. South-west winds blow for much of the year but are stronger and more 
persistant in winter, and are reinforced by thermic wind effects. These raise a rough sea, even 
in the lagoon area behind the barrier reef. In the absence of this wind, a heavy swell of 
remote origin can be detected. Cyclones in the Mozambique channel can also create a 
disturbed sea state for several days. Tides are semi-diurnal, with a maximum range of 3.2 m. 
This is a lower range than occurs in the north of Madagascar, though it is fairly substantial for 
coral reef areas. Low spring tides at noon and midnight have a strong controlling influence on 
the biota of the reef flat; the tidal amplitude is such that there are strong localised currents 
during ebb and flow. Rainfall is sparse in this area, but sufficient rainfall in the interior 
results in permanent rivers and a marked terrigenous component to the sediments near the 
coast. Surface water temperatures vary seasonally from 22.5°C to 27.5°C. Sediments in the 
reef area are described by Thomassin and Cauwet (1985). 

The Bay of Toliara (Tulear) is connected to the sea by two channels, Passe Nord and Passe Sud, 
which dissect Grand Recif, a barrier reef 18 km long and 1100-2900 m wide. Nosy Tafara is 
an islet situated to the south of the Grand Recif. In the southern part of the bay there are 
three lagoon reefs - Beloza, Dimadimatsy, and Norinkazo, which are separated from the shore 
by a littoral channel 1 km wide with maximum depth 2 m. These reefs (about 1000 x 1000 m) 
are separated from each other by passes about 300 m wide and 4-5 m deep, with strong 
currents. North of the inner reefs lie two sandstone banks, Mareana and Ankilibe, which have 
a scattering of isolated coral heads (Clausade et al., 1971). On the south of the bay, there is a 
fringing reef on the west side of the sandy Sarodrana peninsula, 3500 m x 450-1000 m wide 
(Pichon, 1978b). 

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An environmental profile of Madagascar 

REEF STRUCTURE and CORALS There have been numerous ecological and descriptive 
studies of the Toliara reefs (Clausade et al., 1971; Harmelin- Vivien et a!., 1982; Jaubert and 
Vasseur, 1974; Pichon, 1972a, 1972b, 1978a, 1978b; Peyrot-Clausade, 1977; Thomassin, 1978b, 
1983; Thomassin and Cauwet, 1985; Vasseur, 1974, 1977, 1981, 1984; Weydert, 1973a, 1973b). 
Several distinct zones on the Grand Recif have been described in detail by Pichon (1972a, 
1978b). The 'frontorecifal ensemble', or seaward reef slope, is very steep but rarely vertical. 
Scleractinian corals are abundant, extending to at least 30 m deep, depending on local reef 
profiles and sedimentary conditions. The deep seaward slope, 300-400 m wide, gives way to a 
gently sloping terrace which drops from 20 to 50 m, with an average depth of 20 m. The 
morphology of this '20 m terrace' is characterised by large furrows at right angles to the reef, 
the floors of which are sand-filled, with ridges covered with corals. The terrace extends up to 
the edge of the reef flat, which it joins in a short but vertical wall of corals. This wall consists 
of a groove and spur system aligned with the coral ridges on the deeper terrace. The grooves 
are part filled with coral debris; the spurs are constructed of corals and abundant calcareous 
red algae. Numerous tunnels are formed by grooves which have coalesced at the top. Some of 
these caverns communicate with the reef flat by blowholes, while others become blocked with 
sediments. This network of overhangs, grooves and tunnels houses a diverse cryptic 
community of ahermatypic corals and other sessile invertebrates which has been described by 
Jaubert and Vasseur (1974) and Vasseur (1974, 1977). 

The spur and groove structures are best developed on parts of the reef most exposed to the 
south-westerly seas. On the deeper part of the seaward slope below the terrace, coral diversity 
decreases with depth, and there is a simultaneous decline in abundance (Pichon, 1978b). The 
dominant coral family is the Pectiniidae, in particular the genera Echinophyllia 
and Oxypora; Pectinia is less common. Other common genera are the agariciids Leptoseris 
and Pachyseris, a Pocillopora species and the mussids Cynarina lacrymalis and Blastomussa sp. 
Gorgonia and the ahermatypic coral Dendrophyllia are also common, with antipatharians and 
alcyonarians. On the terrace, coral diversity is greater on shallower parts of the terrace and 
species characteristic of the deeper part become less abundant. Common genera are Acropora, 
Porites, Lobophyllia and several faviids. On the buttresses, these species decrease in shallower, 
more turbulent water, and are replaced by encrusting species from the genera Pavona. 
Hydnophora, Montipora and Acropora. The tops of the buttresses support numerous 
Pocilloporidae, and a high cover of calcareous red algae. In the most exposed parts, Acropora 
is the principal genus, but coralline algae may be dominant, although there is no true algal 
ridge either here or in any other part of Madagascar (Pichon, 1972a). 

The vast plateau of the reef flat of Grand Recif is exposed by a few decimetres at low 
spring-tides and has been termed the 'epirecifal ensemble'. On the outer reef flat there is an 
upper platform of spurs, dominated by crustose coralline algae. Shoreward of this is an outer 
moat with algae and corals, followed by an outer pavement with encrusting coralline algae but 
few corals. A boulder tract is then reached, consisting of corals and blocks of limestone 
deposited by strong sea conditions. This provides the highest elevation on the reef flat, 
reaching 1 m above the surrounding level. Several corals, zoanthids and algae grow on the 
boulder surfaces while beneath them is a very rich mollusc, crustacean and echinoderm fauna. 
Inside this there is an inner moat remaining covered at low spring-tides, which is flat and 
covered by sandy patches, with many corals whose growth is truncated by the low water level. 
Microatolls are common, as are groups of corals whose growth is peripheral only. The reef flat 
at this point is friable and irregular. Corals become less abundant as a third zone of seagrass 
beds is reached which lie on a slightly raised part, occupying half of the reef flat. Large 
accumulations of sediments are found here with a typical seagrass-sediment fauna, notably 
echinoderms (Clausade et al., 1971; Pichon, 1978b). 

Behind the Grand Recif, the reef flat gives way to a 'postrecifal ensemble' (Clausade et al., 
1971; Pichon, 1978b), the reef slope descending to a lagoon floor with a maximum depth of 
20 m. Currents mostly run parallel to the shore, and the sediments have an increasing 
terrigenous component as shore is approached. A variety of seagrasses characterize the area, 
with abundant coral patches and pinnacles, particularly near the reef. These are formed largely 
by massive faviid corals, with Acropora, several foliaceous species, and Millepora. Such reefs 



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Marine and coastal ecosystems 

also occur adjoining the mainland, where their tops support seagrasses which grade into the 
littoral zone. 

Altogether, 62 hermatypic coral genera have been reported (Pichon, 1978). Amongst the 
corals, the genus Horastrea appears to be fairly common. Thomassin (1969, 1973, 1978a, 
1978b) and Thomassin et al. (1976) have described the sandy bottom communities of the area. 

NOTEWORTHY FAUNA and FLORA The marine fauna has been extensively studied. For 
example, the fish fauna of the outer slope of the Grand Recif is described by Harmelin-Vivien 
(1977). The mollusc fauna on the boulder tracts is described by Thomassin and Galenon 
(1977); sponge distribution is described by Vacelet and Vasseur (1977). Mangroves are found 
around Saradrano. 

SCIENTIFIC IMPORTANCE and RESEARCH More research has been done on the reef 
morphology and coral communities of the Grand Recif than on any other single reef in the 
Indian Ocean. Accompanying this is a wide range of marine studies with direct relevance to 
the reef and its communities, resulting in a very substantial body of detailed literature. For 
this reason, the reef is of great importance to comparative and temporal studies of reef systems 
in general. The marine station of the University of Madagascar is situated at Toliara. Up until 
1955, the work carried out there was conducted almost exclusively by French researchers 
(generally from the Station Marine d'Endoume, Marseille). Currently, research is carried out 
by Madagascans and visiting scientists. Productivity of the reef has recently been studied by 
Pichon and Morrissey (1985). 

ECONOMIC VALUE and SOCIAL BENEFITS Commercial harvesting of fish and 
invertebrates takes place, the former at least, on an increasingly large scale. The Toliara reefs 
are the main area of exploitation of reef fish, local consumption exceeding 100 tonnes 
annually. Rabesandratana (1985) provides further details. Prawns are collected on the inner 
slope of the lagoon where littoral mangroves grow in proximity to the reefs. The Spiny 
Lobster Panulirus penicillatus is collected for local consumption (see Part V.10). Scylla serrata 
is occasionally caught on seagrass beds on the Grand Recif. Molluscs collected 
include Charonia tritonis, Cypraeacassis rufa, and Pinctada (Meleagrina) margaritifera 
(Rabesandratana, 1984, 1985 and see Part V.8). The massive coral Porites somaliensis is 
collected from the Grand Recif for use in septic tanks and cesspools. Branching corals are also 
collected for sale to tourists in the shell market at Toliara. Corals were exported until at least 
1980, when an export figure of 4.1 tonnes was recorded by the Toliara customs office 
(Rabesandratana, 1985). Shells are exported in large quantities for the ornamental shell trade 
from the Toliara region, and certain species are considered to be coming rare (Rabesandratana, 
1985). The area has a high potential for tourism, as yet largely unrealized. A French-run 
hotel caters for the few SCUBA divers who visit the area and an underwater trail has been set 
up (Pichon, 1983). 

DISTURBANCE or DEFICIENCIES Corals, such as Porites somaliensis, have in ihe past been 
collected for building purposes (Rabesandratana, 1984), though it is unclear to what extent the 
practice continues. Over-fishing is becoming a serious problem. Pichon reported in 1983 that 
up to 200 boats used the Grand Recif daily; in the early 1970s the area was virtually unfished. 
Populations of benthic fishes had been noticeably depleted, though pelagic fish populations 
were as yet apparently largely unaffected (Pichon, 1983). Fishing is carried out with nets, 
often with mesh below the legal size, with harpoons and with toxins extracted from plants 
(e.g. Euphorbia). This last method is noted as being particularly destructive as it kills 
indiscriminately. Amongst invertebrates, Pinctada (Meleagrina) margaritifera has reportedly 
been overharvested to the point of virtual extinction and Cypraeacassis rufa has become 
noticeably rarer; concern has also been expressed for Charonia tritonis, of which large 
numbers, including small specimens, are reportedly on sale in Toliara; minimum size 
regulations, where these exist, are largely ignored (Rabesandratana, 1984). 

LEGAL PROTECTION There is at present no legal protection for the reefs. 



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An environmental profile of Madagascar 

RECOMMENDATIONS The Grand Recif has been proposed as a National Marine Park; this 
designation has been refused twice by the Ministry of Animal Production and Forests 
(Rabesandratana, 1984). The proposal is for the creation of two reserves: an integral reserve, 
containing the Grand Recif de Toliara (within the area 23 o 20 , -23°38 , S, 43 <> 30 , -43 o 42'E), and 
an adjacent partial reserve, containing the 'postrecifal channel', the mangrove area of 
Sarodrano, and the reefs fringing the coastline of Barn-Hill Point. The proposal aims to assure 
conservation of the reef, while developing its potential for scientific research and tourism (both 
national and international) and managing fisheries resources to allow sustained harvests by local 
fishermen. Under the proposed decree, access to the integral reserve will be limited to 
authorised personnel, though this will include tourists accompanied by officials. Licensed 
artisanal fishermen will also be permitted, though fishing will be under strict control (e.g. all 
underwater fishing and use of toxins, explosives and monofilament nets banned). Access to the 
partial reserve will be uncontrolled, though again fishing will be controlled and limited to 
licensed artisanal fishermen. The importance of a local education programme to demonstrate 
the value of such conservation measures is stressed (Rabesandratana, 1984). 



NAME Nosy Be 

GEOGRAPHICAL LOCATION Island off north-west coast; 13°20'S, 48°15'E. 

PHYSICAL FEATURES Nosy Be is a volcanic island on the continental shelf of Madagascar 
(Battistini, 1960). Its coast is a very embayed, low Quaternary plain, and reef flats are found 
around much of the island. Seaward of the reef flats, the reef slopes are extensive, reaching to 
at least 20 m deep, and sometimes to 45 m. Deeper slopes exist, but these are sedimented 
(Pichon, 1971a). 

Dominant seas come from the north-west or north-north-west, while in winter the south-east 
trade winds occur, from which Nosy Be is protected to a great extent by the Madagascan 
mainland. Rain is abundant in the southern summer, and infrequent in the cooler season. In 
common with other reefs near Madagascar, this results in a relatively high proportion of 
terrigenous sediment mixed with the limestone sediments, and turbidity is always high. 
Surface temperatures are relatively stable, ranging from 24° to 29°C. The area is subjected to 
cyclones (UNEP, 1982b), but the swell is usually weak. The reefs experience a tidal amplitude 
of 4.2 m (Pichon, 1972a). 

REEF STRUCTURE and CORALS The geology of the reefs has been described by Battistini 
(1960), and their biology by Pichon (1971a and 1972a). The morphological and 
sedimentological characteristics of Nosy Be, as well as the reef communities, are characteristic 
of a low energy regime. Common corals are fungiids, and crustose coralline algae are lacking. 
Approximately 63 genera have been recorded (Pichon, 1978b). The most typical reefs are 
Andilana Reef on the north-west coast, and Amphoraha and Navetsy Reefs on the north coast. 
These have developed on the most exposed parts of the island and exhibit typical spur and 
groove structures with well developed boulder tracts. On the eastern coast, the reefs 
Antsatrabevoa, Antafianambitry and Befefiky are exposed only to a local breeze which blows 
from the south-east and which therefore has a small fetch. These reefs may have extensive 
reef flats up to 1.5 km wide, a rudimentary spur system, and a small boulder tract both of 
which are thickly covered with corals with typical genera such as Caulastrea. Other anthozoans 
and calcareous red algae are not extensive. On the outer flats is a seagrass bed 
of Thalassodendron ciliatum and Syringodium isoetifolium which forms a 20 m wide, 
uninterrupted strip just in front of the boulder zone, on a layer of sand of skeletal origin 
15-25 cm thick. Seagrass beds are also extensive on the inner parts of the reef flats (Pichon, 
1972a). On the west and south-west, the reefs are poorly developed and not very active; reef 
flats are absent, but their place may be taken by accumulations of broken coral fragments. 

Reef slopes of the fringing reefs extend to only 8 or 10 m depth. A rudimentary spur and 
groove system is discernible only at low tides on the more exposed reefs (it is absent from 
sheltered reefs) and appears to be formed from an alignment of corals rather than substantial 



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Marine and coastal ecosystems 

algal constructions. Coral species are diverse, composed of massive and foliaceous forms, but 
few Acropora species. A notable antipatharian, Eucirripathes, is a constant element on these 
reefs. At 8-10 m depth the rough alignments of corals disappear, and the grooves become 
filled with sediment. On the sedimented slope below the reef slope, communities of the 
corals Heteropsammia michelini and Helerocyathus aequicostatus exist, with some Trachyphyllia 
geoffroyi. In general terms, these reefs are regarded as 'inner reefs' analogous to the inner 
reefs of Toliara to the south (see separate account) (Pichon, 1972a), while the muddy bottom 
coral community is analogous to that of a lagoon floor. In this case, the 'lagoon' extends from 
the base of the fringing reefs out to the reefs of an outer barrier, which is submerged. 

Reef slopes of the outer, submerged barrier formation, which lies to the west, extend deeper 
than those which fringe the island itself. There are two principal types of slope: gentle slopes 
(up to 45°) with rich scleractinian communities, notably tubular Acropora corals to about 45 m 
deep, with abundant Peyssonnelia and Halimeda algae; and near-vertical slopes, which support 
abundant coral to about 20 m, below which diversity is much poorer. At 50-70 m depth, these 
walls are covered with a fine sediment composed of Halimeda fragments. 

NOTEWORTHY FAUNA and FLORA Mangroves are abundant in many of the bays and 
estuaries. Eretmochelys imbricata is reportedly present, though it is not known whether it nests 
here. Cypraea species are abundant and species recorded include C. nucleus, C. diluculum, 
C. onyx, C. oweni, C. lamaki and C. chinensis (Magnier, 1981). The Tropic bird Phaethon 
lepturus lepturus breeds on nearby islands (Cooper et al., 1984). 

ECONOMIC VALUE and SOCIAL BENEFITS Nosy Be is one of the few important tourist 
centres in Madagascar (Jolly, 1980; Magnier, 1981); coral reefs appear to be a significant tourist 
attraction, though it is unclear to what extent their full potential has been developed. Pinctada 
margaritifera has reportedly been heavily exploited, to the point of virtual extinction, having 
formerly been abundant in the area (Rabesandratana, 1984). Ornamental shells are collected in 
the Nosy Be area (Randrianarijaona and Razafimbelo, 1983). 

SCIENTIFIC IMPORTANCE and RESEARCH Previously ORSTOM had a research station at 
Nosy Be which carried out fishery studies. A National Oceanographic Institute is based at 
Nosy Be. Projects have reportedly been funded on fisheries and pollution, though the latter 
does not appear to be a serious problem in this part of Madagascar at present (Pichon, 1983). 

DISTURBANCE or DEFICIENCIES Overexploitation of P. margaritifera (see above). 

LEGAL PROTECTION The small island Nosy Tanikely, about 8 km south of Nosy Be, is 
protected for its terrestrial fauna, and the surrounding waters are considered a Marine Reserve 
but have no legal protection (Rabesandratana, 1984). 

The 740 ha Reserve Naturelle Integrate de Lokobe (R.N.I, no. 6) is situated in the south-east 
corner of Nosy Be. The coast line forms the southern edge of the reserve (See Part VI); Pichon 
(1972a) indicates that at least part of this has a reef front, though it is not clear whether 
protection extends offshore. 



NAME Offshore sand cays: Nosy Foty, Nosy Anambo, Nosy Fasy, Nosy Faty, Nosy Faho, 
Nosy Langna 

GEOGRAPHICAL LOCATION North-west continental shelf; between 12° and 13°20'S, 48 
and 49°E. 

PHYSICAL FEATURES Data for these small islands are taken primarily from Pichon (1972a). 
Low islands with surrounding reef, some (Nosy Fasy and Nosy Faty) emergent only at low 
tide. Nosy Langna also has an outcrop of Cretaceous pre-coralline basement at the level of the 
reef flat. The direction of the swell affecting the reefs is predominantly from the north-west 



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An environmental profile of Madagascar 

and north-north-west, with only gentle seas along the sides facing the Madagascan mainland. 
Temperatures range from about 24°C to 29°C. 

REEF STRUCTURE and CORALS The reefs of Nosy Anambo, Fasy, Faty and Foty represent 
emergent parts of the offshore barrier reef structure (see 'Introduction'). The coral structures 
topped by these sand cays show marked adaptations to the dominant north-west seas. The cays 
generally lie on the leeward edges of the reef flats. Facing the direction of maximum 
exposure, the reef flat is edged by a spur and groove structure which is dominated by 
calcareous red algae, behind which is a boulder tract. Most of the flats have sandy expanses 
alternating with coral formations. On the leeward sides of the cays is a sandy shore, followed 
by large expanses of corals with alcyonarians and Millepora, followed by deeper beds of the 
seagrass Thalassodendron ciliatum (Pichon, 1972a). 



Acknowledgements 

We are very grateful to Dr M. Pichon for assistance with this section. 

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Conference de Madagascar sur la Conservation des Ressources Naturelles au Service du 

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Battistini, R. (1960). Description geomorphologique de Nossi Be, du delta du Sambirano et 

de la baie d'Ampasindava. Mem. I.R.S.M. ser F. III. Pp. 121-143. 
Battistini, R. (1964). Etude geomorphologique de l'extreme sud de Madagascar. Cujas. 

Paris. Pp. 1-536. 
Battistini, R. (1972). Madagascar relief and main types of landscape. In: 

Richard-Vindard, G. and Battistini, R. (Eds). Biogeography and ecology of Madagascar. 

Monogr. biol. Junk, The Hague. Pp. 367-410. 
Clausade, M., Gravier, N., Picard, J., Pichon, M., Thomassin, B., Vasseur, P., Vivien, M. 

and Weydert, P. (1971). Coral reef morphology in the vicinity of Tulear (Madagascar): 

Contribution to a coral reef terminology. Tethys Supp. 2: 74. 
Cooper, J., Williams, A.J. and Britton, P.L. (1984). Distribution, population size and 

conservation of breeding seabirds in the afrotropical region. In: Croxall, J.P., Evans, 

P.G.H. and Schreiber, R.W. (Eds), Status and Conservation of the World's Seabirds. ICBP 

Technical Publication No. 2, Cambridge. 
Daniel, J., (1972). Etude bathymetrique et sedimentologique d'une baie tropicale, la baie 

d'Ambaro. Thesis, Universite de Paris VI. 85 pp. 
Daniel, J., Dupont, J. and Jouannic, C. (1970). Etude de la relation entre le carbone 

organique et l'azote dans les sediments de la Baie d'Ambaro. ORSTOM Nosy Be Doc. 

16: 1-20. 
Daniel, J., Dupont, J. and Jouannic, C. (1972). Relations Madagascar - Archipel des 

Comores (Nord-Est du Canal de Mozambique). Sur la nature volcanique du Banc du 

Leven. C.R. Acad. Sci. Paris 274 D: 1784-1787. 
Fourmanoir, P. (1963). Distribution ecologique des poissons de recifs coralliens et 

d'herbiers de la cote ouest de Madagascar. La Terre et la Vie 1: 81-100. 
Guilcher, A. and Battistini, R. (1967). Madagascar - geographie regionale. 'Les cours 

de Sorbonne', Centre de documentation universitaire, Paris. 
Harmelin-Vivien, M.L., (1977). Ecological distribution of fishes on the outer slope of 

Tulear reef (Madagascar). Proc. 3rd Int. Coral Reef Symp., Miami: 289-295. 
Jaubert, J. and Vasseur, P. (1974). Light measurements: duration aspect and the 

distribution of benthic organisms in an Indian Ocean coral reef, (Tulear, 

Madagascar). Proc. 2nd. Int. Coral Reef Symp.. Brisbane 2: 127-142. 
Jolly, A. (1980). A world like our own - man and nature in Madagascar. Yale University 

Press, New Haven. 



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Marine and coastal ecosystems 

Keiner, A. (1963). Poissons, peche el pisciculture a Madagascar. Centre Technique 

Forestier Tropical, Nogent-sur-Marne. 
Keiner, A. (1965). Contribution a l'etude des eaux suamatres malgaches. Les poissons 

euryhalins et leur role dans le developpement des peches (These Fac. Sciences Paris). Vie 

et Milieu 16(2-c): 1013-1149. 
Keiner, A. (1972). Ecologie, biologie et possibility de mise en valeur des mangroves 

malgaches. Bull. Madagascar 308: 49-84. 
Magnier, Y. (1981). Merveilles sous-marines. In: Oberle, P. (Ed.), Madagascar, un 

sanctuaire de la nature. Lechevalier, Paris. 
ORSTOM (1973). Publications de Centre ORSTOM de Nosy Be. Liste mise a jour au 

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Doc. Sci. Centre ORSTOM Nosy Be 51. 45 pp. 
Pichon, M. (1971a). Comparative study of the main features of some coral reefs of 

Madagascar, La Reunion and Mauritius. In: Stoddart, D.R. and Yonge, CM. 

(Eds) Regional variation in Indian Ocean coral reefs. Symp. Zool. Soc. Lond. 28. 

Academic press, London. Pp. 185-216. 
Pichon, M. (1971b). Horastrea indica n.gen., n.sp., a new hermatypic scleractinian 

coral from the south-west Indian Ocean (Cnidaria, Anthozoa, Madreporaria). Rev. Zool. 

Bot. Afr. 33: 165-172. 
Pichon, M. (1972a). The coral reefs of Madagascar. In: Richard- Vindard, G. r\nd 

Battistini, R. (Eds). Biogeography and ecology of Madagascar. Monogr. biol. Junk, The 

Hague. Pp. 367-410. 
Pichon, M. (1972b). Les peuplements a base de scleractinaires dans les recifs corallians 

de la baie de Tulear (Sud-ouest de Madagascar). Proc. Symp. Corals and Coral Reefs 

(1969). J. mar. biol Ass. India 1972: 173-181. 
Pichon, M. (1974). Free living scleractinian coral communities in the coral reefs of 

Madagascar. Proc. 2nd Int. Coral Reef Symp., Miami. 2: 261-267. 
Pichon, M. (1978a). Recherches sur les peuplements a dominance d'anthozoaires dans les 

recifs coralliens de Tulear (Madagascar). Atoll Research Bull. 222: pp. XXXV, 477. 
Pichon, M. (1978b) Quantitative benthic ecology of Tulear reefs. In: Stoddart, D.R. and 

Johannes, R.E. (Eds). Monographs on Oceanic Methodoloy 5 Coral Reefs: Research 

Methods. UNESCO: 163-174. 
Pichon, M. and Morrissey, J. (1985). Premieres mesures de bilan metabolique dans 

l'ecosysteme recifal de Tulear (Madagascar). C. R. Acad. Sc. Paris 300 Series 3(3):99- 101 . 
Rabesandratana, H.D. (1984). Letter to C. Sheppard / IUCN with enclosures including 

documentation on submission of proposals to establish a Marine Park at Tulear and data 

on collection of molluscs in Madagascar. 
Rabesandratana, H.V. (1985). About some reef utilizations in Madagascar. Proc. 5th Int. 

Coral Reef Cong., Tahiti 6: 661-668. 
Randrianarijaona, P. and Razafimbelo, E. (1983). Rapport national pour Madagascar. Report 

prepared for UNEP Regional Seas East Africa Programme. 
Thomassin, B. (1969). Les peuplements de deux biotopes de sables coralliens sur le Grand 

Recif de Tulear, S.O. de Madagascar. Rec. Trav. Stn. mar. Endoume Suppl. 9: 59-133. 
Thomassin, B. (1973). Peuplements des sables fins sur les pentes internes des recifs 

coralliens de Tulear (S.O. de Madagascar). Essai d'interpretation dynamique des 

peuplements de sables mobiles infralittoraux dans un complexe recifal soumis ou non aux 

influences terrigenes. Tethys Suppl. 5: 157-220. 
Thomassin, B. (1978a). Soft bottom communities. In: Stoddart, D.R. and Johannes, R.E. 

(Eds). Monographs on Oceanographic Methodology 5. Coral Reefs: Research Methods. 

UNESCO: 263-298. 
Thomassin, B. (1978b). Les peuplements des sediments coralliens de la region de Tulear 

(SW de Madagascar). Leur insertion dans le contexte cotier indo-pacifique. Thesis, 

Universite Aix-Marseille II. 494 pp. 
Thomassin, B.A. (1983). Successions of faunistic assemblages in coral reef soft bottom 

biotopes according building evolutional stages in the Tulear region (Madagascar). 15th 

Pac. Sci. Cong. Dunedin Abs. 2: 235-236. 
Thomassin, B.A. and Cauwet, G. (1985). Organic matter distribution in sediments of the 

Tulear coral reef complexes. Proc. 5th Int. Coral Reef Cong.. Tahiti 3: 377-382. 



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An environmental profile of Madagascar 

Thomassin, B. and Galenon, P. (1977). Molluscan assemblages on the boulder tracts of 

Tulear coral reefs (Madagascar). Proc. 3rd Int. Coral Reef Symp., Miami: 113-117. 
Thomassin, B., Vivier, M.H. and Vitiello, P. (1976). Distribution de la meiofaune et de 

la macrofaune des sables coralliens de la retenue d'eau epirecifale du Grand Recif de 

Tulear (Madagascar). J. Exp. Mar. Biol. Ecol. 22: 31-53. 
UNEP (1982a). Environmental problems of the East African region. UNEP Regional Seas 

Reports and Studies No. 12. 
UNEP (1982b). Marine and coastal area development in the East Africa region. UNEP 

Regional Seas Reports and Studies No. 6. 
Vacelet, J. and Vasseur, P. (1977). Sponge distribution in coral reefs and related areas 

in the vicinity of Tulear (Madagascar). Proc. 3rd Int. Coral Reef Symp.. Miami: 113-117. 
Vasseur, P. (1974). The overhangs, tunnels and dark reef galleries of Tulear (Madagascar) 

and their sessile invertebrate communities. Proc. 2nd Int. Coral Reef Symp.. Brisbane 2: 

143-159. 
Vasseur, P. (1977). Cryptic sessile communities in various coral formations on reef flats 

in the vicinity of Tulear (Madagascar). Proc 3rd Int. Coral Reef Symp.. Miami 1: 95-100. 
Vasseur, P. (1981). Recherches sur les peuplements sciaphiles des recifs coralliens de la 

region de Tulear (SW de Madagascar). Thesis, Universite Aix-Marseille II. 348 pp. 
Vasseur, P. (1984). Les peuplements sessiles sciaphiles des recifs coralliens de la 

region de Tulear (SW de Madagascar): resultats synthetiques. Oceanis 10(1): 51-83. 
Verseveldt, J. (1973). Octocorallia from north-western Madagascar. Koninkl. Nederl. 

Akad. Wetensch. Amsterdam Proc. 76. 
Weydert, P. (1973a). Les formations recifales de la region de Tulear (Cote Sur de 

Madagascar). Aperiju de leurs aspects morphologiques, sedimentologiques et de leur 

evolution. Ass. Senegal et Quatern. Ouest Afr. Bull, liaison 37-38: 57-83. 
Weydert, P. (1973b). Morphologie et sedimentologie des formations recifales de la region 

de Tulear, SW de Madagascar. Thesis, Universite Aix-Marseille II. 646 pp. 



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PART V. FAUNA 

The fauna of Madagascar is unique. The large size of the island, its geological history and 
varied climate and topography have resulted in a diverse fauna with a remarkable degree of 
endemicity, both at species and higher taxonomic levels. 

In general, individual faunal groups are less diverse than in equivalent (tropical) continental 
areas - this applies to, for example, birds, mammals, freshwater fish and butterflies (qv); 
however other groups, such as reptiles and terrestrial molluscs, show relatively high species 
diversity, well comparable with continental areas. 

The great majority of native terrestrial species, in all faunal groups, appear to be dependent on 
forested or wooded areas, lending support to the contention that originally (that is before 
human settlement) much of the island was forested - this in contrast to present-day conditions 
whereby only 20-30% of the land area bears woody cover (see Part III. 4). 

The following section provides summaries for: birds; mammals; amphibians and reptiles; 
freshwater fish; lepidoptera; freshwater, terrestrial and marine molluscs; non-marine and 
marine Crustacea; other non-arthropod invertebrates. 

Data sheets for individual species for some of these groups, extracted from the relevant IUCN 
Red Data Book, are provided in Appendix 3 and annotated species lists in Appendix 2. 
Preliminary faunal lists for each reserve are included in Part VI. 

Reference lists are provided at the end of each of the following sections; in addition extensive 
references are provided with the data sheets on individual species in Appendix 3. 

The most important reference work for animal species on Madagascar is the on-going Faune de 
Madagascar (1956- ). Of the 64 volumes published to date, one concerns birds (vol. 35), two 
deal with mammals (vols 36,44), three with reptiles (vols 33,36,47), one with zoogeography 
(vol. 13) and the remainder with invertebrates [in French]. 

The volume Biogeography and ecology in Madagascar (1972, Monographiae biologicae 21, 
edited by R. Battistini and G. Richard-Vindard, Junk Publishers, the Hague), contains chapters 
on the following groups: arachnids, terrestrial molluscs, insects, freshwater and euryhaline fish, 
reptiles, birds, insectivores, rodents, carnivores, and primates [part English, part French]. 

Madagascar, un sanctuaire de la nature (1981, edited by P. Oberle, Lechevalier, Paris) also has 
(more generalized) chapters on invertebrates, reptiles and amphibians, birds, and mammals [in 
French]. 

Key environments: Madagascar (1984, edited by A. Jolly, P. Oberle, and R. Albignac, 
Pergamon Press, Oxford) also has introductory chapters on invertebrates, amphibians, reptiles, 
birds, and mammals with separate chapters for insectivores, carnivores and lemurs (the chapters 
on invertebrates, birds, reptiles and mammals are English translations of the equivalent 
chapters in Madagascar, un sanctuaire de la nature). 



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An environmental profile of Madagascar 



V.l. BIRDS 

The avifauna of Madagascar, as much of the rest of the fauna, is characterised by two factors - 
a relative poverty in number of species (compared with equivalent continental areas), and a 
high degree of endemism at family and lower taxonomic levels. 

Forbes-Watson et al. (1974) listed 250 species in the avifauna of Madagascar, including 2 
introductions, 53 non-breeding visitors, and 197 native breeding species. Of these 197 
residents, 106 are endemic (a further 25 are shared only with the Comoros). Among these 106 
species, there are 32 endemic genera (a further eight are shared only with the Comoros). An 
annotated list of bird species recorded in Madagascar, based on Forbes-Watson et al. (1974) and 
Dee (in press) is provided in Appendix 2. 

The following five families are endemic to the Madagascar Region: 



Mesitornithidae (Mesites) 
Brachypteraciidae (Ground-rollers) 
Leptosomatidae (Cuckoo-roller) 
Philepittidae (Asitys) 
Vangidae (Vangas) 



One subfamily is also endemic to the region: 
Couinae (Cuculidae) (Couas) 



3 spp. 
5 spp. 

1 sp. (also occurs on the Comoros;) 

4 spp. 

14 spp. (one species also occurs on the 
Comoros) 



10 spp. (one probably extinct) 



The Leptosomatidae and Brachypteraciidae have been considered subfamilies of the Coraciidae 
but are now generally recogised as distinct families. 



Status 

One species - Coua delalandei - is regarded as (probably) recently extinct; 27 others are 
currently considered threatened or probably so, 3 all of which are endemic to Madagascar 
(Collar and Stuart, 1985). Four species are considered 'endangered', comprising one 
grebe, Tachybaptus rufolavatus, one duck, Aythya innotata and two raptors, Haliaeetus 
vociferoides and Eutriorchis astur, the second of which is in a monotypic genus. Of the 
remaining species, one is 'vulnerable', twelve are 'rare', five 'indeterminate', and five classified 
as 'insufficiently known'. In addition, a further fourteen species are identified as 
near-threatened, although two of these, Ardeola idae and Circus maillardi, are not endemics. 

The endemic families Mesitornithidae and Brachypteraciidae are of particular note - all three 
species of the former are assigned categories, two as 'rare' (Mesitornis variegata and Monias 
benschi) and one as 'insufficiently known' (Mesitornis unicolor); four of the five ground-rollers 
are classified as 'rare', with Alelornis pittoides being considered near-threatened at present. 

Threatened or possibly threatened bird species in Madagascar are: 



K Tachybaptus pelzelnii 

E Tachybaptus rufolavatus 

K Ardea humbloti 

V Anas bernieri 

E Aythya innotata 

E Haliaeetus vociferoides 

E Eutriorchis astur 



Madagascar Little Grebe 
Alaotra Grebe 
Madagascar Heron 
Madagascar Teal 
Madagascar Pochard 
Madagascar Fish Eagle 
Madagascar Serpent Eagle 



3 Full explanations of threatened species ('RDB') categories are provided in Appendix 3. 

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Fauna 



R Mesitornis variegata 

K Mesitornis unicolor 

R Monias benschi 

I Sarothrura waiersi 

K Amaurornis olivieri 

R Charadrius thoracicus 

Ex Coua delalandei 

I 7yto soumagnei 

R Brachypteracias leptosomus 

R Brachypteracias squamiger 

R Atelornis crossleyi 

R Uratelornis chimaera 

I Neodrepanis hypoxantha 

R Phyllastrephus apperti 

R Phyllastrephus tenebrosus 

R Phyllastrephus cinereiceps 

R Xenopirostris damii 

I Xenopirostris polleni 

K Monticola bensoni 

I Crossleyia xanthophrys 

I Newtonia fanovanae 



White-breasted Mesite 
Brown Mesite 
Subdesert Mesite 
Slender-billed Flufftail 
Sakalava Rail 
Madagascar Plover 
Snail-eating Coua 
Madagascar Red Owl 
Short-legged Ground-roller 
Scaly Ground-roller 
Rufous-headed Ground-roller 
Long-tailed Ground-roller 
Yellow-bellied Sunbird-asity 
Appert's Greenbul 
Dusky Greenbul 
Grey-crowned Greenbul 
Van Dam's Vanga 
Pollen's Vanga 
Benson's Rockthrush 
Madagascar Yellowbrow 
Red-tailed Newtonia 



Full data sheets are provided in Appendix 3 (taken from Collar and Stuart, 1985). 



Geographical distribution 

The geographical distribution of the 106 endemic bird species of Madagascar can be analysed 
on a very simple level by dividing the country into four regions, east, north, south and west, 
corresponding roughly to the major phytogeographic divisions of eastern rainforest, sambirano, 
western deciduous forest and southern thorn bush. 



From this the following figures emerge: 



East 
North 
West 
South 



No. spp. 
in region 

83 (20) 
49(5) 
52(12) 
42(7) 



No. spp. confined 
to region 

30(12) 



2(2) 
10(5) 



Total 106 (28) 

Figures in parentheses indicate number of species assigned RDB categories in each group. 

These figures highlight the over-riding importance of the eastern region of the island in terms 
of the number of species supported - 79% of endemics are found in at least part of the eastern 
region, while 29% are confined there on current knowledge, 40% of which are currently 
considered to be under some degree of threat. The second most important of the four domains 
is the south, which has ten species not known to occur elsewhere on the island - half of these 
are assigned RDB categories at present (Collar and Stuart, 1985). 



REFERENCES 

N.B. An extensive bibliography on the Madagascan avifauna is provided in Appendix 3. 



Benson, C.W. (1981). Les oiseaux: des especes uniques au monde. 
Madagascar, un sancluaire de la nature. Lechevalier, Paris. 



In Oberle, P (ed) 



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An environmental profile of Madagascar 

Collar, N.J. and Stuart, S.N. (1985) Threatened birds of Africa and related islands: 

the 1CBP/IUCN Red Data Book, 3rd edition, part 1. ICBP and IUCN, Cambridge. 
Dee, T.J. (in press). The status and distribution of the endemic birds of Madagascar. 

ICBP, Cambridge. 
Forbes-Watson, A.D., Keith, G.S. and Turner, D.A. (1974). Madagascar bird list. 

Unpublished typescript. 
Milon, P., Petter, J. -J., Randrianasolo, G. (1973). Faune de Madagascar XXXV. Oiseaux. 

ORSTOM and CNRS, Antananarivo and Paris. 



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Fauna 

V.2. MAMMALS 

The living native land mammals of Madagascar are confined to five orders: Primates; 
Chiroptera; Insectivora; Carnivora and Rodentia. A single representative of the order 
Artiodactyla is also present - Potamochoerus larvatus - though this is thought likely to have 
been introduced by man, and there is a recently extinct (subfossil) hippopotamus Hippopotamus 
lemerlei. One member of the order Sirenia - Dugong dugon - occurs in inshore coastal waters. 

The taxonomy of some of these groups, notably the Primates, Insectivores and Rodents, is 
uncertain, and the number of species contained in them is a matter of contention, though the 
great majority are endemic. 

Insectivora The insectivores on Madagascar comprise representatives of two families - the 
Soricidae, of which two widespread species are present (Suncus murinus and Suncus etruscus, 
though the Madagascan form of the latter is sometimes considered a separate, endemic species 
- Suncus madagascariensis) and the Tenrecidae, a family which has been considered endemic 
to the Madagascan region, though the African genera Potamogale and Micropotamogale are 
now generally included, though in a separate subfamily, the Potamogalinae. 

The taxonomy of the Tenrecidae is unstable and many taxa are poorly known, often from only 
one or two specimens; however some 30 species in 9 genera are generally recognized. One 
species Tenrec ecaudatus has been introduced to the Comores, Reunion, Mauritius and the 
Seychelles; all others are confined to Madagascar and its offshore islands. 

Chiroptera Some 28 species of bats have been recorded on Madagascar, nine of these endemic 
(though one, Triaenops humbloti, may only be a colour variant of another, T. rufus), with an 
additional three nearly endemic (one also occurs on the Comores, one on Aldabra and one on 
Reunion). One endemic species Myzopoda durita is in its own monotypic and hence endemic 
family, the Myzopodidae; all other species are in non-endemic genera. 

Primates The number of extant species of Madagascan primates depends on the classification 
adopted. This report recognizes 28 species in four different families - the Cheirogaleidae, 
Lemuridae, Indriidae and Daubentoniidae; the first two families are sometimes lumped together 
in the Lemuridae, while conversely the genera Lepilemur, Hapalemur and Varecia are 
sometimes split off into a separate family, the Lepilemuriaae. The taxonomy of Lepilemur is 
highly complex, with recent classifications varying from a single species with five subspecies to 
seven species, one with five subspecies; in this report seven species are recognized. 

Two species of the genus Lemur are found on the Comores as well as on Madagascar, all other 
species are endemic to Madagascar and its offshore islands. 

Carnivora There are seven indigenous species of carnivore in Madagascar, each in its own 
monotypic genus in the family Viverridae and all endemic. As well as feral dogs and cats, the 
palm civet Viverricula indica has been introduced, though is reported to live predominantly in 
savanna regions near villages and to be generally absent from true forest. 

Rodentia Ten species of native rodents in seven genera are currently recognized on 
Madagascar, all are endemic and are ascribed to the same endemic subfamily, the Nesomyinae, 
of the family Cricetidae. Little is known of the distribution or status of these species: one 
(Eliurus myoxinus) is apparently widespread; two (Hypogeomys antimena and Macrotarsomys 
ingens) have restricted ranges in parts of the west; one {Macrotarsomys bastardi) is widespread 
in the west; the others occur in the eastern forests and have generally been rarely recorded 
though at least some are likely to be fairly widely distributed in this region (Nesomys rufus is 
also known from one specimen collected at Maintirano on the west coast though this individual 
is distinctive and may represent a separate species). 



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An environmental profile of Madagascar 

The following lemurs have been assigned threatened species categories, following the 1986 
IUCN Red List of Threatened Animals: 



K Hapalemur griseus 

E Hapalemur simus 

K Lemur catta 

K Lemur coronatus 

E Lemur macaco flavifrons 

V Lemur macaco macaco 

V Lemur mongoz 

I Lemur rubriventer 

K Lepilemur dorsalis 

K Lepilemur edwardsi 

K Lepilemur leucopus 

K Lepilemur microdon 



K Lepilemur mustelinus 

K Lepilemur ruficaudatus 

K Lepilemur septentrionalis 

I Varecia variegata 

E Allocebus trichotis 

K. Microcebus coquereli 

K Phaner furcifer 

K Avahi laniger 

E Indri indri 

V Propithecus diadema 

K Propithecus verreauxi 

E Daubentonia madagascariensis 



Five of the Madagascan viverrids have been assigned categories as follows: 



V Cryptoprocta ferox 
K Eupleres goudotii 
K Fossa fossa 



K Galidictis fasciata 
K Salanoia concolor 



Insufficient information is available at present to assign categories to mammal species from 
other groups, although amongst insectivores, the aquatic Limnogale mergulus is believed to 
present most cause for concern. 

An annotated list of all non-marine mammal species is given in Appendix 2 and data sheets for 
all lemurs and the Dugong are provided in Appendix 3. 



PRINCIPAL REFERENCES 

N.B. Full references for the lemurs are provided with the data sheets in Appendix 3. 

Albignac, R. (1972) The carnivora of Madagascar. In Battistini, R. and Richard-Vindard, 

G. (eds) Biogeography and ecology in Madagascar. Monographiae biologicae 21. Junk, the 

Hague. 
Albignac, R. (1973) Mammiferes carnivores. Faune de Madagascar 36: 1-209. 
Eisenberg, J.F. and Gould, E. (1984) The insectivores. In Jolly, A., Oberle, P., and 

Albignac, R. (eds) Key Environments - Madagascar. Pergammon Press, Oxford. 
Eisenberg, J.F. and Gould, E. (1970) The tenrecs, a study in mammalian behavior and 

evolution. Smithsonian Contributions to Zoology 27: 1-127. 
Gould, E. and Eisenberg, J.F. (1966) Notes on the biology of the Tenrecidae. Journal of 

Mammalogy 47(4): 660-686. 
Heim de Balsac, H. (1972) Insectivores. In Battistini, R. and Richard-Vindard, G. (eds) 

Biogeography and ecology in Madagascar. Monographiae biologicae 21. Junk, the Hague. 
Honacki, J.H., Kinman, K.E. and Koeppl, J.W. (Eds) (1982) Mammal species of the world. 

Allen Press, Inc. and the Association of Systematics Collections. Lawrence, Kansas, U.S.A. 
Meester,J. and Setzer, H.W. (Eds) (1971) The mammals of Africa. An identification 

manual. Smithsonian Insititution Press, City of Washington. 
Morrison-Scott, T.C.S. (1948) The insectivorous genera Microgale and Nesogale 

(Madagascar). Proceedings of the Zoological Society of London 1 18: 817-822. 
Petter, F. (1972) The rodents of Madagascar: the seven genera of Malagasy rodents. In 

Battistini, R. and Richard-Vindard, G. (eds) Biogeography and ecology in 

Madagascar. Monographiae biologicae 21. Junk, the Hague. 
Petter, J. -J. (1972) Order of primates: sub-order of lemurs. In Battistini, R. and 

Richard-Vindard, G. (eds) Biogeography and ecology in Madagascar. Monographiae 

biologicae 21. Junk, the Hague. 



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Fauna 

Petter, J. -J., Albignac, R. and Rumpler, Y. (1977) Mammiferes lemuriens. Faune de 

Madagascar 4 4: 1-543. 
Schliemann, H. and Maas, B. (1978) Myzopoda aurita. Mammalian species No 116. 

American Society of Mammalogists. 
Tattersall, I. (1982) The primates of Madagascar. Columbia University Press, New York. 
Thomas, O. (1918) On the arrangement of the small Tenrecidae hitherto referred to 

Oryzorictes and Microgale. Annals and Magazine of natural History (9)1: 302-307. 



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An environmental profile of Madagascar 
V.3. AMPHIBIANS AND REPTILES 

The herpetofauna of Madagascar is of very great interest in several respects. 

1. With about 144 amphibians and 257 reptiles, Madagascar is, for its size, relatively rich in 
species (the reptiles are particularly numerous). Many species, including nearly 40 amphibians, 
have been described since 1970; doubtless a number remain to be discovered. Several recent 
descriptions are based on material collected some years ago but only recently studied. 

2. The great majority of species (over 90%) are endemic to the island; this includes all but two 
of the 144 amphibians. 

3. There is a distinct contrast between:- 

a. a small group of species-poor genera, usually endemic and often monospecific, apparently 
relict forms representing archaic lineages present since the first fragmentation of 
Gondwanaland. Examples: Erymnochelys, the boas Acrantophis and Sanzinia, the seven 
malagasy iguanids. 

b. a group of very species-rich genera, some endemic, apparently representing separate 
adaptive radiations from several chance immigrations (eg. by rafting from the African 
mainland) subsequent to the geographic isolation of Madagascar. Examples: Chamaeleo 
and Brookesia among chamaeleons (Madagascar has two-thirds of the world's 
species), Scelotes among scincids, Boophis and Mantidactylus among frogs. 

4. The affinities of the reptile fauna are mainly with Africa, while those of the amphibians 
are with Africa and the Orient. However, the zoogeographic relationships of certain forms are 
especially noteworthy. For example, the pelomedusid turtle Erymnochelys is most closely 
related to the South American genus Podocnemis; similarly the boid genera Acrantophis 
and Sanzinia are most closely related to boas in South America (in both these examples related 
fossil material is known from intervening sites in Africa and/or Europe, but no extant forms 
are present in these areas). The seven malagasy iguanids, very distinct from all other iguanids, 
contribute in large measure to the highly enigmatic distribution of the family (present in 
Madagascar, Fiji-Tonga, and the New World; the family is unknown in Africa). 

5. The microhylid genus Pseudohemisus (a monotypic form endemic to Madagascar) has 
recently been shown to be precisely intermediate in regard to tadpole anatomy between the 
families Ranidae and Microhylidae. The taxonomic position of the Scaphiophryninae, to 
which Pseudohemisus (and Scaphiophryne, tadpole unknown) are assigned therefore remains 
unclear (the possibility that Pseudohemisus is in some sense a link between ranoid and 
microhyloid lineages cannot be discounted since the discovery of 'living fossils' is more likely 
on an island such as Madagascar where ancestral populations, possibly competitively inferior, 
can be isolated from their descendants). 

The taxonomic composition of the herpetofauna is as follows:- 



CLASS: AMPHIBIA 
Family: Hyperoliidae 
Family: Microhylidae 
Subfamily: 

Cophylinae (endemic) 
Dyscophinae (endemic ?) 
Scaphiophryninae (endemic) 
Microhylinae 
Family: Ranidae 
Subfamily: 

Mantellinae (endemic) 
Raninae 
Family: Rhacophoridae 



species (genera) 
8/9(1) 



33 (9) 
3/4(1) 
5(2) 
1(1) 



60(3) 

3(3) 

30(2) 



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Fauna 

CLASS: REPTILIA 

Family: Testudinidae 5 (3) 

Family: Cheloniidae 4 (4) 

Family: Dermochelyidae 1(1) 

Family: Pelomedusidae 4 (3) 

Family: Crocodilidae 1 (1) 

Family: Gekkonidae 63 (12) 

Family: Iguanidae 7 (2) 

Family: Chamaeleontidae 53 (2) 

Family: Scincidae 47 (10) 

Family: Cordylidae 12 (2) 

Family: Typhlopidae 9 (2) 

Family: Boidae 3 (2) 

Family: 'Colubridae' 48 (15) 
(plus 2 sea snakes) 

All the amphibians present are frogs; there are no caecilians, newts or toads. Most of these are 
forest-living treefrogs; there are few savannah forms and very few burrowers. Around 10% of 
the 144 species present can persist or thrive in open and/or human-dominated landscapes; 30% 
live in one of the three high mountain areas; but 60% are restricted to low-medium altitude 
moist forest. A few forest forms appear to be restricted to single localities, but probably most 
are widely distributed in the eastern escarpment rainforest; many are obligate tree-axil dwellers. 

This distribution pattern is not repeated among the reptiles. Presumably due to their much 
greater tolerance of non-humid conditions, many species occur in the seasonal western forests 
and in the truly arid south. 

It has been suggested that the herpetofauna is highly sensitive to human modification of the 
environment; this is presumably of minimal concern to the small number of wide-ranging 
forms abundant in various marginal or secondary habitats, but is likely to be important to the 
many highly localized species and perhaps to rainforest species in general. Examples of the 
former are the ranid frog Ptychadena madagascariensis (occurs throughout, especially in rice 
fields), the hyperoliid Heterixalus betsiteo (common in cleared forest) and the scincid Mabuya 
gravenhorsti (favours secondary 'savoka' vegetation); examples of the latter are the 
tortoise Geochelone yniphora (at Cape Sada), the ranid Mantella aurantica (in the Perinet 
forest), and two Lygodactylus geckos at the summit of Mt Bity. 

Loss of habitat is probably the predominant factor adversely affecting the malagasy 
herpetofauna; other factors include relaxing of traditional tribal taboos ('fady') protecting 
certain forms (eg. tortoises, boid snakes, crocodiles at some localities), and persecution or 
over-exploitation (eg. Nile Crocodile, freshwater turtles). Overall it must be stressed that the 
distribution, ecology and conservation status of most elements of the herpetofauna are 
inadequately known. 

Only one species, the 'Angonoka' Geochelone yniphora, is known to be critically threatened at 
present; it is currently the subject of a conservation project. There are, however, many other 
poorly-known forms of special concern, often known only from a single specimen: for 
example, the colubrid snake Liophidium apperti, known by one specimen collected in 1968 in 
deciduous forest near Befandriana-sud; this forest has now been cleared but for a few isolated 
trees and the survival of the snake must be in question. 

The sea turtles, terrestrial tortoises, the freshwater turtle Erymnochelys madagascariensis, and 
some larger frogs are utilized for food (or trade material, ie. tortoiseshell, in the case of 
Hawksbill Eretmochelys); the condition of these resources should be investigated and 
appropriate management applied. The tortoises Geochelone yniphora and G. radiata, and the 
boid snakes Acrantophis and Sanzinia are nominally protected by law (decree of 16 February, 
1961). Export of all wild animals, or parts thereof, is nominally controlled. 



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An environmental profile of Madagascar 

Some 22 species of amphibians and 70 species of reptiles, listed below have been assigned 
IUCN threatened species categories or are being considered for inclusion in a category. The 
many Insufficiently Known species ('K' in the annotated lists, Appendix 2) do not appear 
below. Those already categorized consist of nine members of the order Testudines 
and Crocodylus niloticus. With these exceptions (marked by an asterisk in the list below), the 
designations are those suggested by the one or two competent authorities, but CMC do not yet 
have the requisite corroborative data, thus these should not be regarded as 'official' IUCN 
designations. In the case of taxa not restricted to Madagascar, the designations refer to the 
world range. 



AMPHIBIA 
Hyperoliidae 

R 'Hyperolius' nossibeensis 
Cophylinae 

R Paracophyla tuberculala 
Dyscophinae 
I Dyscophus antongili 
Mantellinae 

R Laurentomantis horrida 
R Laurentomantis ventrimaculata 
R Mantidactylus argenteus 
R Mantidactylus domerguei 
R Mantidactylus glandulosus 
R Mantidactylus klemmeri 
R Mantidactylus punctatus 
R Mantidactylus webbi 
Rhacophoridae 
R Boophis albilabris 
R Boophis microtis 



R Platypelis milloti 



R Laurentomantis malagasia 

V Mantella aurantica 

V Mantella laevigata 
R Mantidactylus eiselti 

R Mantidactylus grand isonae 

R Mantidactylus peraccae 

R Mantidactylus pseudoasper 



R Boophis leucomaculatus 



Testudinidae 




V* Geochelone radiata 


E' 


I* Pyxis arachnoides 


I* 


Cheloniidae 




V* Caretta caretta 


E = 


E* Eretmochelys imbricata 


E J 


Pelomedusidae 




I* Erymnochelys madagascariensis 




Crocodilidae 




V* Crocodylus niloticus 




Chamaeleonidae 




R Brookesia decaryi 


R 


R Brookesia nasus 


R 


I Chamaeleo antimena 


R 


R Chamaeleo bifidus 


R 


I Chamaeleo campani 


R 


I Chamaeleo fallax 


R 


I Chamaeleo gastrotaenia 


R 


R Chamaeleo mallhe 


R 


I Chamaeleo parsonii 


R 


Scincidae 




R Amphiglossus splendidus 


E 


R Androngo trivittatus 


R 


R Paracontias brocchii 


R 


R Pygomeles braconnieri 


R 



Geochelone yniphora 
Pyxis planicauda 

* Chelonia mydas 

Lepidochelys olivacea 



Brookesia ebenaui 
Brookesia tuberculata 
Chamaeleo balteatus 
Chamaeleo boettgeri 
Chamaeleo cucullatus 
Chamaeleo gallus 
Chamaeleo globifer 
Chamaeleo minor 
Chamaeleo will-si 

Amphiglossus stumpffi 
Mabuya boettgeri 
Paracontias holomelas 
Scelotes ornaticeps 



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Fauna 



Colubridae 

R Alluaudina belly i 

V Geodipsas heimi 

R Heteroliodon torquatus 

V Ithycyphus miniatus 

V Langaha nasuta 

V Liophidium rhodogaster 
?R Liophidium trilineatum 

?E Liopholidophis grandidieri 

R Lycodryas arctifasciatus 

R Lycodryas gaimardi 

R Lycodryas inornatus 

R Lycodryas variabilis 

I Pararhadinea albignaci 

R Pseudoxyrhopus ambreensis 

R Pseudoxyrhopus heterurus 

R Pseudoxyrhopus microps 

V Pseudoxyrhopus tritaeniatus 



R Alluaudina mocquardi 

V Geodipsas infralineata 

V Ithycyphus goudoti 

V Langaha alluaudi 

I Liophidium apperti 

V Liophidium torquatus 
?R Liophidium vaillanti 

I Liopholidophis pinguis 

R Lycodryas betsilineatus 

R Lycodryas guentheri 

R Lycodryas maculatus 

R Micropisthodon ochraceus 

R Pararhadinea melanogaster 

I Pseudoxyrhopus dubius 

R Pseudoxyrhopus imerinae 

R Pseudoxyrhopus occipitalis 

I Pseudoxyrhopus quinquelineatus 



An annotated list of Madagascan Amphibian and Reptile species is provided in Appendix 2, 
and data sheets for those species marked * above in Appendix 3. 



REFERENCES 

Angel, F. (1942). Les Lezards de Madagascar. Afe'm. /Icarf. Malgache. 36. Pp. 1-190, 

Pis. 1-22. (Distribution data for lizards) 
Blanc, Ch.P. (1972). Lei Reptiles de Madagascar et des lies Voisines. Pp 501-614 in 

Battistini, R., and Richard-Vindard, G. (Eds), Biogeography and Ecology in 

Madagascar. The Hague: W. Junk. (Major source of species list). 
Blanc, Ch.P. (1977). Reptiles. Sauriens Iguanidae. Faune de Madagascar, 45, 197 

pp., Paris, Orstom/CNRS. 
Blanc, Ch.P. (1981). Batraciens et Reptiles: formes et couleurs insolites. Chap. 4, pp. 

57-62, in Oberle, P. (Ed), Madagascar, un sanctuaire de la nature. Paris, Lechevalier 

S.A.R.L. 
Blommers-Schlosser, R.M.A. (1979). Biosystematics of the Malagasy frogs. I. Mantellinae 

(Ranidae). Beaufortia No. 352, Vol. 29:1-77. 
Blommers-Schlosser, R.M.A. (1979). Biosystematics of the Malagasy frogs. II. The genus 

Boophis (Rhacophoridae). Bijdragen tot de Dierkunde 49 (2):261-312. 
Blommers-Schlosser, R.M.A. (1982). Observations on the Malagasy frog genus 

Heterixalus Laurent, 1944 (Hyperoliidae). Beaufortia 32(1):1-11. 
Bohme, W., & Meier, Ff. (1980). Revision der madagassischen 

Homopholisf Blaesodactylus) Arten (Sauria: Gekkonidae). Senck. biol. (Frankfurt) 60(5/6): 

303-315. 
Bour, R. (1978). Les tortues actuelles de Madagascar (Republique malgache): liste 

systematique et description de deux sous-especes nouvelles (Reptilia-Testudines). Bull. 

Soc. Et. sci. Anjou, N.S., 10:141-154. 
Brygoo, E.R. (1969). Chamaeleo guentheri Boulenger, 1888, synonyme de C. 

pardalis Cuvier, 1829. Bull. Mus. natn. Hist. nat. Paris, ser 2, 41(1): 119-121. 
Brygoo, E.R. (1971). Reptiles. Sauriens Chamaeleonidae. Genre Chamaeleo. Faune de 

Madagascar, 33, 318 pp., Paris, Orstom/CNRS. 
Brygoo, E.R. (1974). Notes sur les Chamaeleo de Madagascar, XII, Cameleons du 

Marojezy. C. peyrieresi n.sp. et C. gastrotaenia guillaumeti n.subsp. (Reptilia, Squamata, 

Chamaeleonidae). Bull. Acad, malgache 51(1):151-166. 
Brygoo, E.R. (1978). Reptiles. Sauriens Chamaeleonidae. Genre Brookesia et 

complement pour le genre Chamaeleo. Faune de Madagascar, 47, 174 pp., Paris, 

Orstom/CNRS. 
Brygoo, E.R. (1979). Systematique des Lezards Scincides de la region malgache. I. 

Scelotes trivittatus (Boulenger, 1896) nov. comb, synonyme de Scelotes trilineatus Angel, 

1949. Bull. Mus. natn. Hist, nat., Paris, ser. 4, 1, sect. A, N° 4: 1115-1120. 



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An environmental profile of Madagascar 

Brygoo, E.R. (1980). Systematique des Lezards Scincides de la region malgache. II. 

Amphiglossus astrolabi Dumeril et Bibron, 1839; Gongylus pollcni Grandidier, 
1869; Gongylus slumpffi Boettger, 1882 et Scelotes waterloti Angel, 1930. Bull. Mus. nam. 

Hist, nat., Paris, ser. 4, 2, sect. A, N° 2:525-539. 
Brygoo, E.R. (1980). Systematique des Lezards Scincides de la region malgache. III. Les 

"Acontias" de Madagascar: Pseudacontias Barboza du Bocage, 1889, Paracontias Mocquard, 

1894, Pseudacontias Hewitt, 1929, et Malacontias Greer, 1970. IV. Amphiglossus 

reticulatus (Kaudern, 1922) nov. comb., troisieme espece du genre; ses rapports 

avec Amphiglossus waterloti (Angel, 1920). Bull. Mus. natn. Hist. nat.. Paris, 4 e ser., 2, 

1980, section A, n° 3 : 905-918. 
Brygoo, E.R. (1980). Systematique des Lezards Scincides de la region malgache. V. 

Scelotes praeornatus Angel, 1938, synonyme de Scelotes s.l. frontoparietalis (Boulenger, 

1889). Bull. Mus. natn. Hist. nat.. Paris, 4 e ser., 2, 1980, section A, n° 4 : 1155-1 160. 
Brygoo, E.R. (1981). Systematique des Lezards Scincides de la region malgache. VI. 

Deux Scincines nouveaux. Bull. Mus. natn. Hist, nat., Paris, 4 e ser., 3, 1981, section A, n° 

1 : 261-268. 
Brygoo, E.R. (1981). Systematique des Lezards Scincides de la region malgache. VII. 

Revision des genres Voeltzkowia Boettger, 1893, Grandidierina Mocquard, 1894, 

et Cryptoscincus Mocquard, 1894. Bull. Mus. natn. Hist. nat.. Paris, 4 e ser., 3, 1981, 

section A, n° 2: 675-688. 
Brygoo, E.R. (1982). Systematique des Lezards Scincides de la region malgache. IX. 

Nouvelles unites taxinomiques pour les Scelotes s. 1. Bull. Mus. natn. Hist. nat.. Paris, 4 e 

ser., 3, 1981, section A, n° 4: 1193-1204. 
Brygoo, E.R. (1983). Systematique des Lezards Scincides de la region malgache. XI. Les 

Mabuya de Madagascar. Bull. Mus. natn. Hist, nat., Paris, 4th ser., 5, sect. A, No. 4: 

1079-1108. 
Brygoo, E.R. (1984). Systematique des Lezards Scincides de la region malgache. XII. Le 

groupe d'especes Gongylus melanurus Gunther, 1877, G. gastrosiictus O'Shaughnessy, 1879, 

et G. macrocercus Gunther, 1882. Bull. Mus. natn. Hist, nat., Paris, 4th ser., 6, sect. A, 

No. 1: 131-148. 
Brygoo, E.R. (1984). Systematique des Lezards Scincides de la region malgache. XIII. 

Les Amphiglossus du sous-genre madascincus. Bull. Mus. natn. Hist, nat., Paris, 4th ser., 

6, sect. A, No. 2: 527-536. 
Brygoo, E.R. (1984). Systematique des Lezards Scincides de la region malgache. XV. 

Gongylus igneocaudatus A. Grandidier, 1867, et Scelotes intermedius Boettger, 1913. 

Les Amphiglossus du groupe igneocaudatus. Bull. Mus. natn. Hist, nat., Paris, 4th ser., 6, 

sect. A, No. 3: 779-789. 
Brygoo, E.R. (1984). Systematique des Lezards Scincides de la region malgache. XVI. Les 

Amphiglossus du groupe ornaticeps. Bull. Mus. natn. Hist, nat., Paris, 4th ser., 6, sect. A, 

No. 4: 1153-1160. 
Brygoo, E.R., Blanc, C.P., and Domergue, C.A. (1970). Notes sur les Brookesia de 

Madagascar. VII. Brookesia karchei n.sp. du Massif du Marojezy. Annates Univ. Madag. 

(ser. sci.) 7: 267-271. 
Brygoo, E.R., Blanc, C.P., and Domergue, C.A. (1972). Notes sur les Chamaeleo de 

Madagascar. 10. Deux nouveaux cameleons des hauts sommets de Madagascar: C. 

capuroni n.sp. et C. gastrotaenia andringitraensis n.subsp. Bull. Mus. natn. Hist. nat. 

Paris, ser. 2, 42: 601-613. 
Brygoo, E.R., Bourgat, R., and Domergue, C.A. (1972). Notes sur les Chamaeleo de 

Madagascar. C. tuzetae n.sp., nouvelle espece du sud-ouest (Reptilia, Squamata, 

Chamaeleonidae). Bull. Mus. natn. Hist. nat. Paris, ser. 3, 21: 133-140. 
Brygoo, E.R., and Domergue, C.A. (1967(1968)). Description d'un Cameleon nouveau de 

Madagascar, Chamaeleo tsaratananensis n.sp. Bull. Mus. natn. Hist. nat. Paris, ser. 2, 

39(5): 829-832. 
Brygoo, E.R., and Domergue, C.A. (1968). Les Cameleons a rostre impair et rigide de 

l'ouest de Madagascar. Mem. Mus. natn. Hist. nat. Paris, N.S., ser A, 52(2): 1-110. 
Brygoo, E.R., and Domergue, C.A. (1968). Description d'un nouveau Brookesia de 

Madagascar: B. vadoni n.sp. (Chamaeleonides). Bull. Mus. natn. Hist. nat. Paris, ser. 2, 

40(4): 677-682. 



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Fauna 

Brygoo, E.R., and Domergue, C.A. (1969). Un Brookesia des forets orientales de 

Madagascar, B. thieli n.sp. (Chamaeleonides). Bull. Mus. natn. Hist. nat. Paris, ser. 2, 

40(6): 1103-1109. 
Brygoo, E.R., and Domergue, C.A. (1969). Chamaeleo balleatus Dum. et Bib. (dans C. 

et A. Dumeril, 1851) n'est pas synonyme de C. bifidus Brogniart, 1800. Bull. Mus. natn. 

Hist. nat. Paris, ser. 2, 41(1): 104-116. 
Brygoo, E.R., and Domergue, C.A. (1969). Notes sur la Brookesia de Madagascar. IV. 

Une serie de petits Brookesia de Nosy Mangabe (Chamaeleonides). Bull. Mus. natn. Hist. 

nat. Paris, ser. 2, 41(4): 833-841. 
Brygoo, E.R., and Domergue, C.A. (1970). Notes sur les Brookesia de Madagascar. 

Description de deux especes nouvelles: B. lambertoni n.sp. et B. therezieni n.sp. 

(Chamaeleonidae). Bull. Mus. natn. Hist. nat. Paris, ser. 2, 41(5): 1091-1096. 
Brygoo, E.R., and Domergue, C.A. (1970). Notes sur les Chamaeleo de Madagascar. C. 

belalandaensis n.sp., Cameleon du Sud-Ouest. Bull. Mus. natn. Hist. nat. Paris, ser. 2, 

42(2): 305-310. 
Brygoo, E.R., Blanc, C.P., and Domergue, C.A. (1974). Notes sur les Chamaeleo de 

Madagascar, 12. Cameleons du Marojezy. C. peyriersi n.sp. et C. gastrotaenia guillaumeti 

n.subsp. (Reptilia, Squamata, Chamaeleonidae). Bull. Acad, malgache. 51(1): 151-166. 
Busse, K. (1981). Revision der Farbmuster-Variabilitat in der madagassischen Gattung 

Mantella (Salientia: Ranidae). Amphibia-Reptilia 2: 23-42. 
Domergue, C.A. (1983). Notes sur les Serpents de la region malgache III. Descriptions 

de trois especes nouvelles rapportees au genre Liophidium Boulenger, 1896. Bull. Mus. 

natn. Hist. nat.. Paris, ser. 4, 5 (4): 1109-1122. 
Domergue, C.A. (1984). Notes sur les Serpents de la region malgache IV. Descriptions 

d'une espece et d'une sous-espece nouvelles. Bull. Mus. natn. Hist, nat., Paris, ser 4, 

6(1): 149-157. 
Domergue, C.A. (1984). Notes sur les Serpents de la region malgache V. Le genre 

Alluaudina Mocquard, 1894. Bull. Mus. natn. Hist. nat.. Paris, 4th ser., 6., sect. A, No. 2: 

537-549. 
Domergue, Ch.A. (1970). Notes sur les Serpents de la region malgache. Lycodryas 

maculatus (Giinther, 1858), espece des Comores. Description de deux femelles. Bull. Mus. 

natn. Hist. nat. Paris, ser. 2, 42(3): 449-451. 
Domergue, Ch.A. (1972). Etude de trois Serpents malgaches: Liopholidophis lateralis 

(Dum. & Bib.), L. stumpffi (Boettger) et L. thieli n. sp. Bull. Mus. natn. Hist. nat. Paris, 

ser. 3, 103, Zool. 77: 1397-1422. 
Guibe, J. (1958). Les Serpents de Madagascar. Mem. Inst. Sci. Madag. Ser. A, 

XII: 189-260. (Major source for snake distribution data) 
Guibe, J. (1978). Les Batraciens de Madagascar. Bonner Zoologische Monographien 

11, pp. 1-144, plus plates 1-82. 
Pasteur, G. (1967). Note preliminaire sur les Geckos du genre Lygodactylus rapportes 

par Charles Blanc du Mont Ibity (Madagascar). Bull. Mus. natn. Hist. nat. Paris, ser. 2, 

39(3): 439-443. 
Pasteur, G., and Blanc, C.P. (1967). Les Lezards du sous-genre Malgache de Lygodactyles 

Domerguella (Gekkonides). Bull. Soc. Zool. France, 92 (3): 583-597. 
Pasteur, G., and Blanc, C.P. (1973). Nouvelles etudes sur les Lygodactyles (Sauriens 

Gekkonides), I. Donnees recentes sur Domerguella et sur ses rapports avec la 

phytogeographie Malgache. Bull. Soc. Zool. France 98(1): 165-174. 



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An environmental profile of Madagascar 

V.4. FISHES 

This account has largely been drawn from Kiener (1963), Kiener and Richard- Vindard (1972), 
Moreau (1979,1983a, 1983b,1984), and FAO (1980,1983). 

The native freshwater fish fauna of Madagascar is relatively depauperate, presumably in part 
as a result of the early geographical isolation of the island. It consists largely of representatives 
of euryhaline groups and contains few of the characteristic major families inhabiting the 
freshwaters of Africa or Indo-Malaya. However, a high proportion of the freshwater fish 
species are endemic, also probably as a result of the early isolation of Madagascar; these 
include all the freshwater atherinids and the nine native cichlids, some of which may be among 
the most primitive in the family. Some species, such as Oxylapia polli and Rheocloides 
Pellegrini, are endemic to isolated river basins within the island, apparently thus restricted by 
physical barriers, often the saline water at the mouth of the rivers. This is particularly so on 
the eastern side of the island where the steep river profiles, narrow coastal plain and low tidal 
amplitude give rise to sharp salinity gradients at the river mouths. On the west coast river 
profiles are much shallower with a far broader coastal plain and continental shelf, and much 
higher tidal amplitude; salinity changes in rivers and estuaries are correspondingly more 
gradual, favouring the development of a euryhaline fauna. Habitat diversity is generally 
greater along this coast, with extensive mangrove and coral reef development (see part IV). 
There is also a greater chance of colonization from African continental waters in the west. 

Information on the present status and distribution of the native fish fauna is scanty and further 
research is desirable. Given the size of the island and the variety of aquatic biotopes, it is 
possible that several species remain as yet undiscovered (T. Roberts, pers. comm.). 

Threats 

Given the sparsity of recent information on the fish fauna, it is difficult to discuss the manner 
in which it is threatened other than in a general and speculative way. 

The increasing human population has led to increasing pressure on the land and, in particular, 
to a decline in the extent of the forest cover, principally through clearance for 'tavy' 
(slash-and-burn) cultivation (see Part III. 3). In turn, this is likely to have had serious effects 
on the fish fauna as the forests exert significant control over the water regime in their 
catchment areas. Thus, for example, deforestation tends to lead to an increase in the volume of 
water passing to rivers as runoff; large discharges are liable to remove soil from denuded banks 
leading to an increase in turbidity and loss of habitat for many species. The Betsiboka River 
has been cited in this regard, as have many high altitude and forest streams which now become 
turbid following only slight rainfall. Waterborne silt may cover the substrate and aquatic 
vegetation, thus reducing the availability of food. Water temperature may rise when tree cover 
is removed (particularly in the headwaters of a catchment), and this may affect the lifecycle of 
both fish and aquatic invertebrates by reducing the viability of eggs and the survival of the 
young. 

Introduced species, both animal and plant, also pose problems. The water hyacinth (Eichhomia 
crassipes) has invaded many lakes and rivers in Madagascar and has blocked several of them. 
Consequent reduction in water flow can lead to increased siltation and reduction of visibility; 
the latter is particularly likely to affect active predatory species which hunt by sight. 
Introduced fish species, several of which are now of great economic importance, have almost 
certainly had considerable impact on the native fish fauna, either as competitors or predators. 
These include Oreochromis spp. which are predominantly herbivorous and have high 
reproductive rates; these could have adverse effects on habitats by reducing vegetation cover 
and thus removing breeding and feeding sites for fish, invertebrates and birds. Paretroplus 
kieneri, for example, typically found in heavily vegetated areas, could be so affected. 

Few native Madagascan fish are currently of interest to the aquarium trade, an exception 
being Bedolia geayi. However, difficulties in transporting fish may have hindered 
development of the trade. Overall, inland fisheries effort appears to have declined in recent 
years, although over-fishing has almost certainly occurred in easily accessible sites; however 

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Fauna 

most fishing effort appears to be aimed at introduced species and no native species are thought 
to be threatened with extinction as a direct result of fishing. 

Threatened species 

The following species have been identified as likely to be threatened by Kiener (1983). His 
categories should be regarded as provisional and not as official IUCN designations. 

'Highly threatened' Oxylapia polli 

Ptychochromoides betsileanus 

'Rare' Acentrogobius therezieni 

Oryzias madagascariensis 
Pachypanchax playfairi 
Rheocloides pellegrini 
Typhleotris madagascariensis 
Typhleotris pauliani 

'Vulnerable' Paratilapia polleni 

Paretroplus dami 
Paretroplus kieneri 
Paretroplus maculatus 
Paretroplus petiti 
Ptychochromis oligacanthus 

An annotated list of all endemic species is included in Appendix 2. 



FISHERIES 

Inland 

The inland fisheries of Madagascar account for 80% of total fish harvest. Approximately 
550 000 ha of the 600 000 ha of waterbodies present on the island may be potentially 
exploitable for fisheries (Kiener and Richard- Vindard, 1972). Freshwater fisheries research is 
carried out by the Division des Recherches Piscicoles (Department of Fisheries Research) 
which has two principal research stations supported by a number of secondary units. 

The principal species caught at present are introduced tilapias and carp. This is in contrast to 
former years when the native Paratilapia polleni was most important. Principal target groups 
in order of importance are: 

1. Cichlidae (primarily introduced tilapias since 1950, although prior to this native cichlids 
were important). 

2. Mugilidae (both marine and freshwater). 

3. Cyprinidae (an introduced group). 

4. Anguillidae (most widely distributed group in the island). 

In addition the Ariidae, Chandidae, Gobiidae and Eleotridae are all important. 

Aquaculture: 25 000 ha of rice fields are now used for fish production; there are also ca 1000 
ha of freshwater fish ponds (FAO, 1983). Formerly, over 85 000 ponds existed, mainly 
concentrated around Antananarivo and Fianaratsoa; this total is now nearer to 10 000 
(Moreau, 1984). Aquaculture could be substantially increased, perhaps by 480 000 ha, by 
including further rice fields and mangrove areas, although the construction of hatcheries is 
likely to be a constraint to future development. Carp and tilapia are the most widely used 
fishes and Chanos chanos has been successfully cultured in brackish water. A project, financed 
by FAO/UNEP, designed to assist with inland fisheries and aquaculture ended in 1982. 

Fishing: As in many African countries, inland fishing tends to be a part-time activity 
undertaken when time is available away from other agricultural activities. The small size of 
the canoes used has resulted in a concentration of activity near to the lake shores where fish 

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An environmental profile of Madagascar 

stocks (particularly juveniles) are now being overexploited. This problem is thought to be 
particularly serious in the Pangalanes, Lake Alaotra, Lake Itasy, lower Betsiboka River, and 
around Toamasina, Antananarivo, and Taolanaro, where there are large numbers of fishermen. 
Elsewhere there is a high potential for fishing, but few fishermen. Moreau (1983a) considers 
that there has been a gradual change from commercial fishery towards subsistence fisheries, 
largely as a result of difficulties with transport and obtaining and repairing equipment. Fewer 
fish are now reaching the large markets which are apparently experiencing shortages. Fish 
consumption is reportedly declining overall. Most fish (75%) is consumed fresh; salting is 
rarely used for preservation and fish are generally smoked if they are to be taken any distance 
for sale. 

Production: Production figures in 1972 are as follows (Kiener and Richard- Vindard, 1972): 
25 000 tons/annum (25 400 tonnes/annum) in natural waters (45 kg/ha/year) 
2000 tons/annum (2032 tonnes/annum) in fishponds (1 tonne/ha/year) 
160 tons/annum (162 tonnes/annum) in rice fields (400 kg/ha/year) 
Total 27 160 tons (27 594 tonnes)per annum. 

These differ from the FAO catch statistics for 1975-1980, where the annual catch for 
cyprinids, cichlids and other freshwater species was estimated at 13 000, 23 000 and 5000 
tonnes respectively, totalling 41 000 tonnes for each year. These estimates will be approximate 
at best: collection of fisheries statistics is extremely difficult as fishing is a very widespread, 
generally low level activity, largely for local subsistence. 

The following observations on inland fisheries in different parts of Madagascar have been 
located. 

River Fisheries: These are generally for subsistence rather than for commercial purposes and 
are poorly quantified. Recruitment in certain areas may be poor, and breeding that normally 
occurs in flooded areas will not take place in drought years. The introduction of trout at sites 
over 1700 m altitude does not appear to have benefitted river fisheries. Fishing in rivers used 
to occur on a larger scale than at present; more use is currently made of artificial water bodies 
and lakes (Moreau, 1983b). 

North West: Commercial eel fisheries are important here, particularly for Anguilla 
mossambica. In the lower part of the Betsiboka River, tilapia and carp had a potential fishery 
of around 2000 tonnes/year, but the catch today is not known. There is little information for 
the remainder of the region (Moreau, 1983b). 

Pangalanes: these artificial channels could be highly productive for a large variety of fishes. 
However, in 1979, with relatively low fishing effort, production was only ca 8 kg/ha. 
Migrating fish, such as the mugilids, are present; some, such as Liza macrolepis, have declined 
(Moreau, 1983b). 

Masianka lagoon: This 15 sq. km lagoon, south of the Pangalanes, has a potential production of 
140 tonnes. However, there is currently low fishing effort soley for subsistence use (Moreau, 
1983b). 

Taolanaro: A series of separate lagoons of 2500 ha has been providing a catch of around 30 
tonnes annually with relatively low fishing effort (Moreau, 1983b). 

Lake Alaotra: P. polleni, C. carpio and tilapia are the major species caught. Eels are also 
taken. In 1963-1967 a peak catch of around 3000 tonnes per annum was reached, decreasing 
by 1976 to approximately 2000 tonnes. This was a similar figure to those of the mid-1950s 
although the fishing effort had greatly increased. Catches have probably since decreased 
(Moreau, 1983b). 

Lake Kinkony: Projected maximum sustainable yield is ca 700 tonnes per annum; possibly half 
of this is currently being caught, principally tilapia. The lake is too far from Antananarivo for 
easy marketing (Moreau, 1983b). 



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Fauna 

Lake Ihotry: Projected maximum sustainable yield is 150 tonnes per annum. There is, 
however, no fishery as the use of fishing equipement is proscribed by a 'fady' (Moreau, 1983b). 

Lake Itasy: Ptychochromoides betsileanus has declined due to the introduction of the water 
hyacinth. Oreochromis niloticus is the main fish caught at present. However, recruitment has 
substantially declined and catches decreased to 275 tonnes in 1976. This lowered catch may 
also be attributed to difficulties in obtaining new nets and other equipment (Moreau, 1983b). 

Marine 

Traditional subsistence marine fisheries are limited to coastal lagoons and shallow inshore 
waters protected by coral reefs. Some 80% of the artisanal fleet works off the west coast. The 
east coast and north and south extremities of the island are difficult to work due to poor 
weather, currents and difficult bottom substrates. The artisanal marine catch is about 
8000 tons/year (8,128 tonnes). The boats are simple wooden outrigger canoes, 4-8 m long, 
propelled by oars or sails. Vessels only carry one or at the most two fishermen using hook and 
line, gill nets, beach seines or traps. Approximately half the marine catch is marketed fresh 
locally in the towns, absence of a distribution system generally preventing internal marketing 
of smoked, dried or frozen marine fish. Freshwater fishes are cheaper than marine fishes 
which are usually more expensive than meat. Experimental fishing in the early seventies 
encouraged industrial exploitation by foreign enterprise and in 1974, 10 000 tons (10 160 
tonnes) of skipjack tuna were landed. Agreements with Japanese, Russian and East German 
fishing interests have been reached; the resultant fishing is thought to have seriously depleted 
the stocks. International fishing has now ceased due to internal problems and the international 
economic crisis. The marine waters are not thought to be particularly rich in fish resources, 
the total available potential (excluding tuna) being about 150 000 tonnes per year, although not 
all of this would be economically exploitable. The most promising demersal species are the 
Sparidae and the Lutjanidae. However, catch rates in exploratory fishing have not been high. 
Areas where pelagic shoaling fishes may be found appear to be highly localised, with very few 
on the western side south of 16°S. North of this, anchovy and sardinella shoals appear to be 
more frequent. FAO/UNDP assisted the marine sector until 1974. 

A guide to the commercial fishes of Madagascar has been produced by FAO (Bauchot and 
Bianchi, 1984). 



REFERENCES 

Arnoult, J and Bauchot, R. (1963). Compte rendu de Mission de Madagascar. Bull. Mus. 

Nat. d'Hist. nat. 1963: 219-227. 
Arnoult, J. (1959). Poissons des eaux douces. Faune de Madagascar IRSM, 

Tananarive 10: 169 pp. 
Arnoult, J. (1963). Un oryziine (Pisces, Cyprinodontidae) nouveau de Test de 

Madagascar. Bull. Mus. Nat. d'Hist. nat. 2e Ser. 35(3): 235-237. 
Bardin, T. (1983). A la recherche des cichlides endemiques de Madagascar. Revue 

Francaise des Cichlidophiles 29: 21-34. 
Bauchot, M.L. and Bianchi, G. (1984). Guides des poissons commerciaux de Madagascar 

(especes marines et d'eaux saumatres). Fiches FAO d'identification des especes pour les 

besoins de la peche. FAO, Rome. 135 pp. 
Boulenger, G.A. (1909-1916). Catalogue of the freshwater fishes of Africa in the 

British Museum. British Museum (Natural History), London 4 vol. 
Daget, J. and Moreau, J. (1981). Hybridation Introgressive entre deux especes de 

Sarotherodon (Pisces: Cichlidae) dans un lac de Madagascar. Bull. Mus. Nat. d'Hist. nat. 4e 

Ser. A. Zool. 3(2): 689-703. 
FAO (1980). Yearbook of fishery statistics. Vol.50. FAO Rome. 
FAO (1983). Fishery country profile. FID/CP/MAG/Rev 3. 4 pp. 
Gunther, A. (1859-1870). Catalogue of fishes in the British Museum. 8 vol. British 

Museum. 



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An environmental profile of Madagascar 

Hoese, D.F. and Winterbottom, R. (1979). A new species of Lioteres, Pisces, Gobiidae 

from Kwazulu with a revised checklist of South African Gobies and comments on the 

generic relationships and endemism of west island ocean goboids. Royal Ontario Museum 

Life Sciences Occasional Paper 31: 1-13. 
Kiener, A. (1960-1961). Poissons malgaches. Liste de noms malgaches de poissons d'eau 

douce, d'eaux saumatres et d'especes euryhalines. Bull. Madag. mai et juin. Au total 117 

pp. 
Kiener, A. (1960-1961). Afrique - Madagascar sous le signe des grands lacs (Mission 

Rhodesie, aout 1960). Bull. Madag. dec. et janv. 175-176: 52 pp. 
Kiener, A. (1963). Poissons, peche et pisciculture a Madagascar. Centre Technique 

Forestier Tropical Pub 24. Nogent sur Marne. 245 pp. 
Kiener, A. (1964). Gobioidei (Pisces) nouveaux ou rares a Madagascar. Bull. Mus. 

Nat. d'Hist. nat. 35(4): 328-333 
Kiener, A. (1966). Contribution a la biogeographie de quelques especes ichtyologiques 

malgaches. Compte Rendu Sommaire des Seances de la Societe de Biogeographies). 

373-374: 3-18. 
Kiener, A. (1983). In litt. 31-3-83. 
Kiener, A. and Mauge, M. (1966). Contributions a l'etude systematique et ecologique des 

poissons cichlidae endemiques de Madagascar. Mem. Mus. Nat. d'Hist. nat. Ser A. Zool. 

49: 49-99. 
Kiener, A. and Richard-Vindard, G. (1972). Fishes of the continental waters of Madagascar. 

pp 477-499. In Biogeography and Ecology in Madagascar. (Eds) Battistini, R. and 

Richard-Vindard, G. Monographic Biologicae 21. W. Junk, The Hague. 765 pp. 
Losse, G.F. (1968). A new record of the anchovy Stolephorus heterolobus (Ruppell) 

from the coast of Madagascar. Cah. ORSTOM ser Oceanogr. 6(2): 117-119. 
Mauge, A.L. (1984). Diagnoses preliminaires d'Eleotridae des eaux douces de Madagascar. 

Cybium 8(4): 98-101. 
Moreau, J. (1979). Le lac Alaotra a Madagascar: cinquante and d'amenagement des peches. 

Cah. ORSTOM ser Hydrobiol 8(3-4): 171-179. 
Moreau, J. (1983a). In litt. 9 September. 
Moreau, J. (1983b). Pers comm. 
Moreau, J. (1984). In litt., 13 December. 
Pellegrin, J. (1914). Sur une derniere collection de poissons recueillie a Madagascar par 

feu F. Geay. Bull. Mus. Nat. d'Hist. nat. 1914(3): 111-113. 
Pellegrin, J. (1914). Sur une collection de poissons de Madagascar. Bull. Soc. 

zool. France 39: 221-234. 
Pellegrin, J. (1919). Sur les Eleotris des eaux douces de Madagascar, description 

d'une espece nouvelle. Bull. Soc. zool. France 44: 266-271. 
Pellegrin, J. (1932a). Poissons de Madagascar recueillis par M. Decary. Description 

d'une variete nouvelle. Bull, de la Societe Zoologique de France 57: 291-297. 
Pellegrin, J. (1932b). Poissons nouveaux de Madagascar recueillis par M. Catala. Bull. 

Soc. zool. France 57: 424- 425. 
Pellegrin, J. (1932c). Poissons de Madagascar recueillis par M. Waterlot. Description 

d'une variete nouvelle. Bull, de la Societe Zoologique de France 58: 225-228. 
Pellegrin, J. (1933). Les poissons des eaux douces de Madagascar et des iles voisins. 

Mem. Academie Malgache. Tananarive, 14: 224 pp. 
Pellegrin, J. (1934). La faune ichtyologique des eaux douces de Madagascar. Paris Ann. 

Sci. Nat. Zool. 18: 425-432. 
Pellegrin, J. (1935). Poissons de Madagascar recueillis par M. Catala, description d'un 

Sicydium nouveau. Bull. Soc. zool. France 60: 69-73. 
Roberts, T. (1981). In litt. to A. Wheeler. 1/10/81. 
Sauvage, H. (1891). Histoire Naturelle des Poissons. Collection Histoire Physique, 

Naturelle, Politique de Madagascar. Impr. Nationale. 543 pp. 
Smith, J.L.B. (1965). Fishes of the family Atherinidae of the Red Sea and the Western 

Indian Ocean with a new freshwater genus and species from Madagascar. Rhodes Univ. 

Dept. Ichthyology, Ichthyol. Bull. 31: 601-632. 
Smith, M.M. (1983). Common and scientific names of fishes of S. Africa. Pt 1. 

Marine Fishes. JLB Institute of Ichthyology, Rhodes University Special Publication 14. 

178 pp. 



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Fauna 

V.5 LEPIDOPTERA: RHOPALOCERA (BUTTERFLIES) 

The island of Madagascar is separated from mainland Africa by straits which are nowhere less 
than 400 km wide. Madagascar has been separated from the mainland for at least 60 million 
years and probably much longer (Owen, 1971). The island is rich in flowering plants but 
relatively poor in animals, especially when compared with other large islands like New 
Guinea. Nevertheless, there are some striking endemic groups of animals, including butterflies 
(Owen, 1971). The Malagasy subregion became isolated before any major evolutionary 
development of butterflies had occurred, and probably before butterflies had properly diverged 
from their ancestors. The ancestors of the present Malagasy fauna must therefore have flown 
or travelled in other ways across the ocean, most of them probably from Africa (Owen, 1971). 
Colonization from the distant African mainland must occur infrequently and rather randomly, 
and this is presumably why the Malagasy butterfly fauna is rather poor in species but at the 
same time rich in endemics (Owen, 1971). 

The biogeographical affinities of the butterfly families are strongly Afrotropical, as is the case 
for most Malagasy arthropods. For the Papilionidae at least, dispersion to Madagascar seems to 
have been from eastern rather than southern Africa. Hancock (1982) states that there is no 
evidence of dispersal from Madagascar back to the African mainland, but Owen (1971) notes 
that Graphium evomber of Madagascar is replaced by a very similar species, Graphium junodi, 
in a narrow strip along the African coast facing Madagascar. 

There are some important exceptions to the generalization that the Malagasy fauna has its 
origins in Africa. Atrophaneura antenor, a swallowtail in the otherwise Oriental and 
Neotropical tribe Troidini, appears to have evolved from an Indian ancestor before that genus 
reached Asia proper (Hancock, 1982). The genus Euploea (Danaidae) is essentially Oriental and 
Australasian in distribution and, although not found on Madagascar, E. euphon is endemic to 
the Mascarenes, and E. mitra to the Seychelles. 

A chain of mountains runs from north to south in Madagascar; the eastern side is wet and 
forested, the western side is relatively dry with a variety of woodland and savanna ecosystems. 
Species richness and endemicity tend to be high in the highland and forested areas, and low in 
lowland and more arid regions. However, for some butterfly families forests are not the 
preferred habitats (e.g. many Hesperiidae, Lycaenidae and Pieridae). 

The southern, and particularly south-western, regions are rather poor in species while the 
deciduous and evergreen forests of the western, northern and eastern regions are richer. Some 
Papilionidae are confined to the deciduous western forests (Papilio morondavana, P. 
grosesmithi), which are clearly important ecosystems for protection. Other species and genera 
are confined to the rain forests (e.g. Graphium endochus (Papilionidae) and Charaxes species 
(Nymphalidae)). The central highlands and eastern rain forests probably include the most 
important localities of all. High altitude areas in central and northern Madagascar include 
important localities, such as Montagne d'Ambre and the Massif de Tsaratanana. 

The butterfly fauna is strongly endemic at the generic, specific and subspecific levels, although 
many other lesser-known insect groups exhibit even higher levels. There are over 300 species 
of butterflies distributed in over 80 genera in the Malagasy subregion (Madagascar, Comoros, 
Mascarenes and Seychelles), of which 233 are endemic (Owen, 1971). At least 17 genera are 
endemic to Madagascar (see Table 6). The levels of species endemism in Madagascar itself are: 
Papilionidae 77% (10 out of 13 species), Pieridae 34% (10 out of 29 species), Nymphalidae 78% 
(133 out of 170 species), Libytheidae 50% (1 out of 2 species), Riodinidae 100% (3 species) and 
Lycaenidae 58% (25 out of 43 species). In addition, there are many endemic subspecies within 
these families. There are relatively few Lycaenidae in Madagascar because the Lipteninae are 
absent. The Hesperiidae have not been considered here but are well represented in the area. 
Seven of the seventeen genera endemic to the Malagasy subregion are in the Hesperiidae 
(Owen, 1971). 

An annotated list of Butterfly species, in all families except the Hesperidae, occurring in 
Madagascar is included in Appendix 2. The conservation status of the Papilionidae, 



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An environmental profile of Madagascar 

Charaxidinae, Acraeinae, Danainae and Nymphalidae has been assessed, but it has not yet been 
possible to assess the threats to the Pieridae, Satyrinae, Libytheidae, Riodinidae or Lycaenidae. 



Conclusions 

The main findings are summarized in Tables 6 and 7. If the relative numbers of species 
identified as threatened in the families considered are extrapolated to the entire butterfly fauna 
of Madagascar, then about 45-50 species may be assumed to be threatened. 

Those species in the families Papilionidae and Nymphalidae identified as threatened in 
Madagascar are: 

R Graphium endochus R Papilio grosesmithi* 

V Papilio morondavana* R Papilio mangoura* 

R Euxanthe madagascariensis R Charaxes cowani 

R Neptis decaryi I Neptis sextilla 

R Apaturopsis kilusa R Smerina manoro 

R Acraea sambavae R Neptis metella 1 

1 Panafrican but very scarce in Madagascar. 

* Data sheets for these species are provided in Appendix 3; notes for all species considered are 
given in Appendix 2. 



TABLE 6. ENDEMIC BUTTERFLIES FROM THE MALAGASY SUB-REGION 

Genus Family Number of species 

Genera in which all species are Malagasy sub region endemics 

Gideona Pieridae 1 

Smerina Nymphalidae: Nymphalinae 1 

Houlbertia Nymphalidae: Satyrinae 8 

Masoura " 5 

Admiratio " 1 

Heteropsis " 2 

Strabena " 41 

Saribia Riodinidae 3 

Trichiolaus Lycaenidae 2 

Rysops " 1 

Hovala Hesperiidae 5 

Fulda " 4 

Amelia " 3 

Malaza " 3 

Miraja " 9 

Perrotia " 6 

Ploetzia " 1 

Other genera with five or more Malagasy sub region endemic species 

Papilio Papilionidae 7 

Charaxes Charaxidinae 8 

Acraea Acraeinae 13 

Henotesia Satyrinae 45 

Hemiolaus Lycaenidae 5 

Lepidochrysops " 5 

(from D'Abrera, 1980; Owen, 1971; Viette, 1956) 

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Fauna 



TABLE 7. THREATENED MADAGASCAN BUTTERFLIES 



Family 


Number of 


Madagascar 


Subregional 


Nui 


mber 


% of total 




species 


endemics 


endemics 


threatened 


threatened 


Papilionidae 


13 


10 


1 




4 


31 


Pieridae 


29 


10 


5 




? 


? 


Nymphalidae 














Danainae 


3 


1 


1 




1 


33 


Charaxidinae 


9 


8 







2 


22 


Nymphalinae 


41 


17 


4 




4 


10 


Acraeinae 


17 


9 


4 




2 


12 


Satyrinae 


100 


98 


1 




? 


? 


Libytheidae 


2 


1 







? 


? 


Riodinidae 


3 


3 







? 


? 


Lycaenidae 


43 


25 


3 




? 


? 



Totals 



260 



182 



19 



(13) 



REFERENCES 

D'Abrera, B. (1980). Butterflies of the Afrotropical Region. Lansdowne Editions, 

Melbourne. 593 pp. 
Hancock, D.L. (1982). Classification of the Papilionidae (Lepidoptera): a phylogenetic 

approach. Smithersia 2: 1-48. 
Owen, D.F. (1971). Tropical Butterflies. Clarendon Press, Oxford. 
Paulian, R. (1951). Papillons Communs de Madagascar. L'Institut de Recherche 

Scientifique, Tananarive-Tsimbazaza. 90 pp. 
Paulian, R. (1956). Insectes Lepidopteres Danaidae Nymphalidae, Acraeidae. Faune de 

Madagascar 2, 102 pp. Publ. Inst Rech. Scient., Tananarive-Tsimbazaza. 
Paulian, R. and Viette, P. (1968). Insectes Lepidopteres Papilionidae. Faune de 

Madagascar 27, 97 pp., 19 pi. ORSTOM, CNRS, Paris. 
Viette, P. (1956). Insectes Lepidopteres Hesperiidae. Faune de Madagascar 3, 83 pp. 

Publ. Inst Rech. Scient., Tananarive-Tsimbazaza. 



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An environmental profile of Madagascar 

V.6. TERRESTRIAL MOLLUSCS 

Since 1949, Professor Fischer-Piette has been studying the snails of Madagascar at the Museum 
National d'Histoire Naturelle Paris, where there is now a collection of some 10 000 lots, 
comprising what is probably the most important Madagascan shell collection. The collection 
consists mainly of shells with a few preserved specimens. More than 30 papers were published 
during 1949-1977 by Fischer-Piette. New species continue to be described from this collection 
(Tillier, 1979). In 1980, van Bruggen published a preliminary checklist of the terrestrial 
molluscs of Madagascar, which has been used as a basis for the accompanying list (van 
Bruggen, 1980b). The fauna has been recently revised by Professor Fischer-Piette, and the 
revision will be published in the Faune de Madagascar (Fischer-Piette et al., in press). 

Madagascar has a land snail fauna currently considered to consist of about 380 species (130 
prosobranchs and 248 pulmonates, but future revisions will probably change these figures 
considerably), which is considered one of the most interesting in the world. Twenty-five 
families (6 prosobranch, 19 pulmonate) and 56 genera (15 prosobranch, 41 pulmonate) are 
known. There are 361 endemic species (127 prosobranch, 234 pulmonate), 11 endemic genera 
(4 prosobranch, 7 pulmonate) and no endemic families. The country has a relatively large 
number of terrestrial prosobranchs (34% of the species present, 27% of the genera, 24% of the 
families) compared to neighbouring South Africa, and the rate of endemism in this group is 
particularly high (35% of all species and 36% of all genera) (van Bruggen, 1980b). The 
dominant families are entirely different from those found in Africa, where the Streptaxidae, 
Urocyclidae, Subulinidae and Achatinidae are most important. Although all these families 
occur on Madagascar, they are generally sparingly represented, and the dominant families are 
Cyclophoridae, Pomatiasidae, Acavidae and Ariophantidae. Accounts of the biogeography of 
the fauna can be found in Fischer-Piette and Blanc (1972), and Verdcourt (1972) mentions the 
fauna in the context of East Africa. 

Not surprisingly, since these are easiest to collect, the best known genera are those containing 
large species, such as Tropidophora, Ampelita, Helicophanta and Clavator. It is to be expected 
that further survey work will considerably alter the relative importance of different families 
and genera, as the smaller species found in habitats such as forest soils become better known 
(Fischer-Piette and Salvat, 1972; Fischer-Piette and Blanc, 1972). Species in the north tend to 
have the flattest shells, and those in the south the most pointed shells. Many of the rain forest 
species are slug-like, and new species are certainly still awaiting discovery. At least 82 (23%) 
of the endemic species are known only from their type localities. 

In general, the calcareous areas of the north, west and south have the richest mollusc fauna. 
Little is known of the ecology of the Madagascan malacofauna but usually terrestrial snails 
prefer a humid habitat. The great majority of species which have been collected come from 
the primary forests, particularly in the north. Using the sparse data given in the original 
descriptions, it has been calculated that of the 142 endemics for which some form of habitat 
description is given, 91 (64%) species are forest dwellers, 34 (24%) are associated with caves, 
and 27 come from other habitats. Many of those for which there are no habitat data can be 
deduced to be forest dwellers by reason of the collection locality. Most species are found along 
or not far from the coast. Tropidophora and many other genera are not found in the centre of 
the island, possibly because these species are not adapted to grassland habitats (Fischer-Piette, 
1948). Millot (Fischer-Piette, 1947) commented that snails were not even found in primary 
forest in the centre of the island (e.g. foret de la Mandraka, foret de Manjakatompo on the 
slopes of Ankaratra, woods of Ambohimanga, forests of Marovatro and Ankaroaka near Lake 
Alaotra), although some surveys have shown that at least some species are found there (see 
Appendix 2). 

The northern quarter of the island, the most diverse in terms of climate, physical and 
geological features and vegetation, is strikingly important for molluscs, with the greatest 
number of endemic snails, and over 56% of known species, occurring there. The Massif de 
Tsaratanana is of major interest, with a very particular fauna including many species in the 
endemic genera Ampelita and Acroptychia (Fischer-Piette, 1952). Montagne d'Ambre exhibits 
a remarkable microendemism for plants, probably associated with the climate and physical 



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Fauna 

features of the area, and probably linked to the mollusc endemism (comment by Heim in 
Fischer-Piette, 1947). 

Certain species survive in habitats ostensibly unsuitable for molluscs. Species of the 
genus Clavator, which is found in the southern part of Madagascar, remain under several 
centimetres of sand in dry weather. After rain they appear one or two hours later, sometimes 
very abundantly, to feed among bushes (Fischer-Piette and Salvat, 1963). Other species have 
an unusual relationship with spiders. The spider Olios coenobita, found in the bush on the 
plateau of Mahafaly, uses snail shells for shelter. Empty shells, sometimes 20 times the weight 
of the spider, are carried up into the bushes and attached by silk threads (Griveaud 
1981). Ampelita chlorozona, for example, was described from a shell collected hanging from a 
bush in Beloha. The Snail-eating Coua Coua delalandei, known from and likely to have been 
endemic to Nosy Borah but probably now extinct (see Appendix 3. A), fed principally on snails; 
it is not known which species were preferred but they may have been Achatinidae rather than 
any of the endemics. 



Endemic genera 

(Number in parentheses indicates approximate number of species) 

Acroptychia (11) 

Ampelita (65) 

Contains many abundant and large species. At least 37 of the species have very localised 
distributions (about 17 in the north); one is known to be widespread. Used to be eaten in 
large quantities by the people of Lake Alaotra (Griveaud, 1981). 

Bat hi a (1) 

Boucardius (8, probably about 40 in new revision) 

Clavator (12) 

This genus, with elongated shells, is one of the most characteristic of the island. It is known 
as a fossil from Africa and has been used for geological dating (Griveaud, 1981). 

Helicophanta (15) 

Includes several species with large and attractive shells. 

Kalidos (52) 

At least 25 species are very localised (13 in the north), 3 are more dispersed and 4 are 
widespread. 

Leucotaenius (7) 

Species in this genus have medium-sized, unusually shaped shells, elongated and heavy. 

Madecataulus (2) 

Malarinia (1) 

Malagarion (1) 

The following genus, although not endemic, is of particular importance in Madagascar: 

Tropidophora 

89 species, of which 87 are endemic. 

The largest genus on Madagascar, containing species with comparatively large shells, 
exceeding 2 cm. The genus Tropidophora is also found in the Comoros, Seychelles, and 
south-east Africa, but has undergone its main radiation in Madagascar where species are 
most numerous, largest and most varied. Almost all the species found there are endemic and 

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An environmental profile of Madagascar 

very few are found throughout the island. The species found on Madagascar have been 
extensively studied and more is probably known about them than about any other genus. They 
have been divided into a number of groups on the basis of their shape (Fischer-Piette and 
Millot, 1949): 

T. cuvierana group (includes occlusa, deliciosa): shells have two very distinct keels. 

T. deshaysiana group (includes moulinsii, vittata, virgo): shells are sculpted and very flattened. 

T. aspera group (includes fulvescens): globose, spiral stripes. 

T. semidecussata group (includes macareae, pyrostoma). 

T. sikorae group (includes filostriata, balteata): shells are elongated. 

T. philippiana group (includes coquandiana): shells are conical and smooth, aperture small. 

T. formosa group (includes pulchella, deburghiae, reticulata): shells are very variable. 

T. tricarinata group: many varieties, although tricarinala itself is rare. 

T. lineata group (includes goudotiana, consocia, vesconis. johnsoni, virgata): shells are small, 

group least well known as little collected. 

Some species, e.g. T. tricarinata, show remarkable polymorphism, thought to be linked to 
habitat. At least 55 species are highly localised (30 in the north), at least ten of the others 
are more dispersed and a further ten, at least, are very widespread. Their distributions fall 
into four main groups (Fischer-Piette, 1947,1948), which fit in well with those areas 
established by Perrier de la Bathie for climate and vegetation (see Part III. 2): 

a. East from Midongy, south of Ambohivoangy. A narrow, uniform band where T. 
bicarinata is dominant. Tropical humid climate, and relatively abundant primary forest. 
Due to the similar climate on Nosy Be and Nosy Komba, these islands have similar fauna. 
Two magnificent species, formosa and deburghiae, are found in the Mananara region at 
Foulpointe and Fenoarivo Atsinanana (Fischer-Piette, 1948). 

b. South and south-west from Taolanaro to north of Toliara. T. philippiana and T. 
coquandiana are dominant (also balteata). These are species with shells with the highest 
spires. Subdesert. 

c. West, from Toliara to Ambongo and Sambirano. T. macareae is dominant. Long dry 
season, calcareous soil; most of forest destroyed. 

d. A small area in the extreme north including Port Leven, Antseranana, Montagne d'Ambre, 
Ankara and Nosy Be. No one species is dominant but the highest number of species are 
found there. Many of the species have very flattened shells ranging from the 
giant cuvieriana to the tiny lamarcki, and very localised distributions (Fischer-Piette, 1948). 



Threats to molluscs 

a. Habitat destruction 

Humidity is particularly important to molluscs which is why primary rain forest tends to be a 
favourite habitat. Opening up of the forest exposes snails to the danger of desiccation, 
particularly as many of the species are slug-like and cannot retract completely into their shells. 
This is discussed in more detail in the IUCN Invertebrate Red Data Book (Wells el al., 1983), 
where several examples are given of species in other parts of the world which have become 
extinct or are now endangered as a result of loss of their forest habitat. It has become clear 
that much of the Madagascan forest has already been lost or is in the process of disappearing 
through a variety of factors (see Part III. 4); this is undoubtedly the main threat to most 
Madagascan land snails. 

b. Introduced species 

A major threat to terrestrial molluscs in the Pacific has been the introduction of the 
carnivorous snail Euglandina rosea from Florida to control the Giant African Snail Achalina 
fulica (Wells el al. 1983). In most cases, E. rosea has had little effect on A. fulica, often 

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Fauna 

preferring to feed on smaller native snails. In Madagascar, A. fulica is a serious pest, especially 
in vanilla plantations on the East coast (Ranaivosa, 1971), and in the Sambirano region where it 
is reported to attack cocoa trees (Griveaud, 1981). Although regarded as a delicacy in many 
countries, it is apparently not eaten by the local people in Madagascar. Attempts at biological 
control by introduced carnivorous snails were tried on a number of occasions: 



1. Euglandina rosea 
January 1965: 



5 snails introduced at Station de l'lvoloina (Taomasina); 
laid eggs and six months later six specimens were 
recaptured at various localities in the station. 



November 1966-December 1968: 



738 adults and 497 eggs at Ivoloina. 

158 adults and 20 eggs at Antalaha (Station de la 

Vanille). 

Most of these came from breeding colonies established 

at Antananarivo, 250 only having been imported from 

Mauritius (where they had been introduced from 

Hawaii and Bermuda). 



March 1970: 



122 from Trinidad released at Ivoloina. 



Releases were always carried out in sites which were regularly frequented by A. fulica. Eggs 
were released after a period of incubation in a laboratory. In 1971, specimens of E. rosea were 
found regularly at the station and empty shells of A. fulica were being found increasingly 
regularly. It was considered that these introductions were being successful. 

2. Gonaxis quadrilateralis and G. kibweziensis 



March 1965: 



110 specimens of G. quadrilateralis 

21 specimens of G. kibweziensis 

Both species introduced from Hawaii; none survived. 



October 1969: 



120 specimens of Gonaxis sp. were raised and laid eggs, 
but by March 1970 only about 40 individuals still 
survived; these were sent to Grande Comoro for release. 



From these trials it was concluded that Gonaxis were not as good biological control agents 
as Euglandina. 

3. Edentulina ovoid ea Brugiere 

A carnivorous species, endemic to Mayotte, which is known to be an active predator 
of Achat ina. 



March 1970: 



120 specimens released at Ivoloina. 



Further introductions were planned. 

Ranaivosa could draw no conclusions in his 1971 paper on the long-term success of these 
introductions, but was hopeful that E. rosea and Edentulina ovoidea would prove useful. 
Long-term studies on the introduced populations were envisaged. There is no information 
available on the outcome of these studies and it is quite possible that the introductions have 
died out, but it is most important that the danger of further introductions is made clear; these 
are detailed in Tillier and Clarke (1983). Surveys should be carried out to determine whether 
any of the introduced populations have survived. 



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An environmental profile of Madagascar 

c. Collecting 

Land snails are known locally as 'Akora', and are not collected very much by the Madagascans. 
However, many species have extremely attractive shells, comparable to those from the 
Philippines, Papua New Guinea and Florida which are currently involved in the shell trade. 
For example, Ampelita viridis is particularly attractive, bright to olive-green in colour, with 
brown stripes and 2-3 cm high. The genus Clavator contains a number of attractive, large 
(over 6 cm long), yellowish-brown shells. The genus Tropidophora contains many species with 
attractively shaped, striped (although not particularly colourful) shells, notably the large T. 
cuvierana with its two keels. Other species reach up to 10 cm in size. Although the 
attractiveness of these species may be a useful tool in conservation publicity, it is 
recommended that firm measures are taken to ensure that commercial collecting does not start. 
Interest in terrestrial species among shell collectors has grown markedly in recent years and 
there would be a ready market for many of the Madagascan species. Since many of these have 
apparently extremely limited distributions and are still very poorly known to science, collection 
should currently be limited to specimens for scientific purposes. 



Recommendations 

a. Further research 

Almost nothing is known of the ecology of most of the Madagascan mollusc fauna and there is 
urgent need to determine this for some of the more important groups in order to make 
recommendations for their conservation (Fischer-Piette and Salvat, 1972). Since most of the 
work has been carried out on collections of shells, there is certainly a need for examination of 
whole preserved specimens for verification and clarification of some of the taxonomical 
problems. Although most parts of the island have been studied, the faunas of the south-east 
(between Ihosy-Betroka-Antaniniria and the Indian Ocean), and the area north of the lower 
Mangoky river are still poorly known (Paulian 1983). 

b. Management of reserves 

Many of the reserves appear to have extensive mollusc faunas (see Part VI. 2). It has been 
estimated that about 160 of the endemic species may occur in reserves. All the reserves should 
be inventoried and efforts should be made to determine which species receive sufficient 
protection from the current system of reserves and which warrant further attention. Mount 
Tsaratanana requires special attention since it is the most important site for molluscs. 

c. Introductions and commercial exploitation 

See above. 

We are very grateful to Professor E. Fischer, Mme. D. Vukadinovic, S. Tillier, P. Bouchet, B. 
Salvat, C. Blanc, B. Verdcourt and A.C. van Bruggen, for their assistance with this section. 



REFERENCES 

Bruggen, A.C. van (1967). Miscellaneous notes on southern African Gastropoda. Euthyneura 

(Mollusca). Zool. Verh. Leiden 91: 1-34. 
Bruggen, A.C. van (1980a). A preliminary checklist of the terrestrial molluscs of 

Madagascar. Achatina 8: 147-164. 
Bruggen, A.C. van (1980b). Notes on the African element among the terrestrial molluscs of 

Madagascar. Haliotis 10(2): 32. 
Decary, R. (1950). La Faune Malgache, son role dans les Croyances et les usages 

indigenes. Payot, Paris. 236 pp. 
Dupouy, J. (1966). Les Veronicellidae de Madagascar, des Comores, des Seychelles, de la 

Reunion et de Pile Maurice. Bull. Mus. Natn. Hist. Nat. Paris (2)37: 667-677. 



-84- 



Fauna 

Dupouy, J. (1966). Les Veronicellidae de Madagascar. Une espece nouvelle: Vaginula 

(Imerinia) fischerii. Bull. Mus. Natn. Hist. Nat. Paris (2)37: 1013-1023. 
Etienne, J. (1973). Inst. Recher. Agronom. Trop. Cult. Vivr., Reunion, Rapport Annuel 

1973: 71-73. 
Fischer-Piette, E. (1945). Recolte malacologique du Professeur Humbert dans le nord de 

Madagascar. Bull. Mus. Natn. Hist. Nat. Paris (2) 17(1): 41-46. 
Fischer-Piette, E. (1947). Distribution des especes du genre Tropidophora (Moll. 

Gast.) a Madagascar. C.R. Somm. Soc. Biogeogr. 24(205): 21-22. 
Fischer-Piette, E. (1948). Premier essai de definition de districts malacologiques 

terrestres a Madagascar. Mem. Inst. Sci. Mad. Ser. A, 2: 161-167. 
Fischer-Piette, E. (1949). Mollusques terrestres de Madagascar Genre Trodiphora (= 

Tropidophora). J. Conchyl. 89: 5,41,111. 
Fischer-Piette, E. (1950). Au sujet de la repartition de Pupoides coenogrictus 

Hutton. J. Conchyl. 90: 221. 
Fischer-Piette, E. (1950). Mollusques terrestres de Madagascar. Genre Helicophanta. 

J. Conchyl. 90: 82-106. 
Fischer-Piette, E. (1952). Mollusques terrestres de Madagascar. Genre Ampelita. 

J. Conchyl. 92: 5-60. 
Fischer-Piette, E. (1963). Mollusques terrestres de Madagascar. Genre Leucotaenius. 

J. Conchyl. 103: 15-23. 
Fischer-Piette, E. (1964). Mollusques terrestres de Madagascar. Famille Enidae. J. 

Conchyl. 104: 67-76. 
Fischer-Piette, E. (1965). Mise au point de nomenclature. Bull. Mus. Natn. Hist. Nat. 

Paris 37(5): 818. 
Fischer-Piette, E. and Bedoucha, J. (1964). Mollusques terrestres de Madagascar Famille 

Streptaxidae. Bull. Mus. Natn. Hist. Nat. Paris 36(2): 368-376. 
Fischer-Piette, E. and Bedoucha, J. (1965). Mollusques terrestres de Madagascar. Famille 

Vertiginidae. Bull. Mus. Natn. Hist. Nat. Paris 37(2): 145-150. 
Fischer-Piette and Bedoucha, J. (1965). Mollusques terrestres opercules de Madagascar. 

Mem. Mus. Natn. Hist. Nat. Paris 33: 50-91. 
Fischer-Piette, E., Bedoucha, J. and Salvat, F. (1966). Mollusques terrestres de 

Madagascar. Ariophantidae. Mem. Mus. Natn. Hist. Nat. Paris 40: 1-50. 
Fischer-Piette, E. and Blanc, F. (1972). Le peuplement de mollusques terrestres de 

Madagascar. In: Battistini, R. and Richard-Vindard, G. (Eds), Biogeography and ecology 

of Madasgascar. Monographiae Biologicae 21. Junk, the Hague. 
Fischer-Piette, E., Blanc, F. and Salvat, F. (1969). Complement aux mollusques opercules 

terrestres de Madagascar. Mem. Mus. Natn. Hist. Nat. Paris (55): 109-144. 
Fischer-Piette, E., Blanc, F. and Salvat, F. (1975). Complement aux mollusques pulmones 

de Madagascar. Bull. Mus. Natn. Hist. Nat. Paris (3)288 (Zool. 198): 209-276. 
Fischer-Piette, E., Blanc, C, Blanc, F., Salvat, B., Salvat, F. and Vukadinovic, D. (in 

press). Mollusques terrestres de Madagascar. 2 volumes: I. Opercules, II Pulmones. Faune 

de Madagascar. 
Fischer-Piette, E., Blanc, F. and Vukadinovic, D. (1974). Additions aux mollusques 

terrestres de Madagascar. Bull. Mus. Natn. Hist. Nat. Paris (3)218 (Zool. 146): 465-524. 
Fischer-Piette, E., Cauquin, M. and Testud, A.M. (1973). Mollusques terrestres recoltes 

par M. Soula dans la region d'Antalaha (Madagascar). Bull. Mus. Natn. Hist. Nat. Paris 

(3)122 (Zool. 94): 477-530. 
Fischer-Piette, E. and Garreau de Loubresse, N. (1964). Mollusques terrestres de 

Madagascar Famille Acavidae. J. Conchyl. 104(4): 129-160. 
Fischer-Piette, E. and Garreau de Loubresse, N. (1964). Mollusques terrestres de 

Madagascar. Genre Propebloyetia. J. Conchyl. 104(4): 161-168. 
Fischer-Piette, E. and Millot, J. (1949). Iconographie et classification des principaux 

Tropidophora de Madagascar (Mollusques Cyclostomidae). Mem. Inst. Sci. Mad. A 

3(1): 57-64. 
Fischer-Piette, E. and Salvat, F. (1963). Mollusques terrestres de Madagascar. Genre 

Clavator. J. Conchyl. 103(2): 53-74. 
Fischer-Piette, E. and Salvat, F. (1972). Originalite des mollusques terrestres de 

Madagascar. In: Comptes rendus de la Conference Internationale sur la conservation de la 

nature et de ses resources d Madagascar, Tananarive. 7-11 October. IUCN Publ. N.S. 

Suppl. Doc. 36. 

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An environmental profile of Madagascar 

Fischer-Piette, E. and Testud, A.M. (1964). Mollusques terrestres de Madagascar 

Achatinaceae. Bull. Mus. Natn. Hist. Nat. Paris (2)36: 377-383. 
Fischer-Piette, E. and Vukadinovic, D. (1971). Sur des mollusques terrestres de 

Madagascar. J. de Conchyl. 109: 29-30. 
Griveaud, P. (1981). Les invertebres: un univers prodigieux. Ch. 3. In Oberle, P. (Ed.), 

Madagascar: un sanctuaire de la nature. Lechevalier, Paris. 
Paulian de Felice, L. (1950). Oniscoides nouveaux de Madagascar. Naturaliste 

Malgache 2: 101-106. 
Paulian, R. (1983). In litt. 28 July. 
Petit, G. (1935). Enumeration systematique des mollusques terrestres et fluviatiles. In: 

Germain, L., Etude de la reserve naturelle du lac Manampetsa. Ann. Sci. nat. Zool. 

18: 421-481. 
Ranaivosoa, H. (1971). Sur deux operations de lutte biologique entreprises par 1'I.R.A.M. 

dans l'archipel des Comores. Agr. Trop. 26: 341-347. 
Salvat, F. (1967). Cyathopoma pauliani n.sp. Mollusque opercule terrestre de 

Madagascar. Bull. Mus. Natn. Hist. Nat. Paris (2)39: 965-967. 
Testud, A.M. (1965). Sur la distribution de Vitrina madagascariensis Smith. J. de 

Conchyl. 105: 230-231. 
Tillier, S. (1979). Malagrion paenelimax gen. nov., spec, nov., a new slug-like 

helicarionid from Madagascar (Pulmonata: Helicarionidae). The Veliger 21(3): 361-368. 
Tillier, S. and Clarke, B.C. (1983). Lutte biologique et destruction du patrimoine 

genetique: le cas des mollusques Gasteropodes Pulmones dans les Territoires fran?ais du 

Pacifique. Genetique, Selection, Evolution 15(4): 559-566. 
Van Goethem, J. (1977). Revision systematique des Urocyclinae. (Mollusca, Pulmonata, 

Urocyclidae). Ann. Mus. Roy. Afr. Centr. Tervuren 8 (218). 
Verdcourt, B. (1972). The zoogeography of the non-marine Mollusca of East Africa. J. 

Conch. 27: 291-348. 
Wells, S.M., Pyle, R.M., and Collins, N.M. (1983). IUCN Invertebrate Red Data Book. 

Conservation Monitoring Centre, Cambridge. 



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Fauna 

V.7. FRESHWATER MOLLUSCS 

Twenty nine species of freshwater gastropod have been described in Madagascar including 21 
prosobranchs and 8 pulmonate basommatophorans. Five species have been described from 
brackish- water or damp biotopes. Checklists are given in Brygoo (1968a), Fischer-Piette and 
Vukadinovic (1973), and Starmuhlner (1969). The total fauna is twice as large as that of 
Kenya, a country similar in area to Madagascar, but the level of endemicity is much lower than 
in the Madagascan terrestrial molluscs. Cleopatra and Melanatria are endemic genera; 
two Afrogyrus species, Bulinus obtusispira and B. bavayi are endemic species. Lake Alaotra, 
the largest lake in Madagascar apparently lacks any endemics. The fauna probably contains 
species derived from the original Gondwanaland fauna, and also species which have reached 
the island since its formation (Brown, 1980). 

Starmuhlner (1969) relates the fauna to three geographical and climatic regions: 

1. The freshwaters of the central steppe-like subtropical highlands have no prosobranchs 
except Pila cecillei but have all the pulmonates except Bulinus mariei, including Afrogyrus 
which is found only in streams above 1800 m. 

2. The escarpments which descend steeply to the east and north-west coasts with extensive 
primary forest have soft water streams and rivers where Melanatria and Cleopatra are 
common. Cleopatra species dominate in the smaller brooks with weaker flow 
whereas Melanatria, a primitive genus, is typical of strongly flowing rivers of the coastal 
tropical rain forest belt. 

3. In the south-east and western areas, the surface waters are richer in dissolved salts. A 
total of nine species has been recorded from the lower Mangoky river (Degremont, 1973), of 
which the two commonest were Lanistes grasseti and 'Anisus' (probably Afrogyrus 
crassilabrum). Melanoides tuberculatus and Radix hovarum may also be dominant. The 
coastal sections of streams have the neritids and Thiaria amarula, with Cerithidia decollata 
dominating in the brackish water zone (Starmuhlner, 1969; Brown, 1980). 



Conservation 

No information is available on the conservation status of endemic freshwater snails. Species 
occurring in or near reserves are listed in Part VI.2. Since most of the endemic species, and 
particularly the genera Cleopatra and Melanatria, are restricted to upland forested areas it is 
probable that they are highly vulnerable to the current extensive deforestation. The acquisition 
of further data relating to the conservation status of these species should be considered a high 
priority. 



Schistosomiasis 

Schistosomiasis or bilharzia is considered briefly here because of its close relationship with 
some of the endemic freshwater snails. Its role in Madagascar is the subject of a large number 
of publications, produced mainly by the Institut Pasteur in Madagascar (see bibliography of 
publications of Inst. Pasteur Mad.). Some of the more important ones are given in the 
references. Brygoo (1968a) provides a detailed survey of the infection in Madagascar, and 
more recent discussions of the Situation are found in Degremont (1973), Brygoo (1972) and 
Brown (1980). 

Two forms of bilharzia or schistosomiasis are found in Madagascar. Intestinal bilharzia is 
caused by the parasitic trematode Schistosoma mansoni, and has Biomphalaria pfeifferi 
(considered a separate species, B. madagascariensis, by Starmuhlner, 1969) as its intermediate 
host. It prevails in the east and on the plateaux; B. pfeifferi appears to be uncommon in 
western Madagascar (Brygoo, 1972; Pfluger, 1977). Its ecology and biology are described in 
Pfluger (1977a,b). In eastern Madagascar it occurs in various habitats including irrigation 
channels and ricefields (Brown, 1980). 

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An environmental profile of Madagascar 

Urinary bilharzia, which prevails in the west, is caused by 5. haematobium, and is carried by 
the endemic snail Bulinus obtusispira which has a high infection rate (Brygoo, 1972; Wright, 
1971). Unlike Biomphalaria, Bulinus does not tolerate temperatures lower than 4°C for longer 
than 24 hours. Bulinus obtusispira is most closely related to B. africanus but was probably 
isolated long ago on Madagascar. It was once thought to be synonymous with B. liratus, a 
species which may be a recently arrived isolate of B. tropicus from South Africa. B. obtusispira 
and liratus are found together in the lower Mangoky, where they have different ecological 
requirements, but their relative distributions elsewhere remain to be worked out (Brown, 
1980). B. liratus is probably unimportant in natural transmission as it has proved resistant to 5. 
haematobium in the laboratory (Wright, 1971), although two snails from the lower Mangoky 
area are reported to have shed cercariae (Degremont, 1973). It is, however, an intermediate 
host for Paramphistomum microbothrium (Prod'hon et al., 1968). 

The different temperature requirements of Bulinus and Biomphalaria are largely responsible for 
the general distribution of schistosomiasis throughout Madagascar. It is predominantly an 
infection of rural and agricultural communities and is increasing in many countries as a result 
of the development of water resources and the modernisation of agricultural methods. The 
creation of man-made lakes, the introduction of new irrigation schemes or the extension of 
existing ones are important factors in the spread of infection since all these activities provide 
suitable habitat for snail vectors (Brygoo, 1972; Webbe, 1981). Ideal habitat is often created 
through careless engineering or construction work. 

Control of schistosomiasis is the subject of intensive research in many countries. Recent 
progress is reviewed in Christie (1978), Hoffman et al. (1979) and Webbe and Jordan (1982) 
and it is concluded that elimination of the disease, rather than eradication of infection, should 
be the goal. A combination of control methods is likely to be most effective, but the 
over-riding need in most countries is the development of low cost methods. 

Chemical control, using molluscicides, has been used extensively and is likely to remain 
important, although it needs to be made more cost-effective. The majority of synthetic 
molluscicides are very expensive and all appear to be toxic in some way to the environment, 
particularly to fish. There is increasing interest in the possibility of using plants with 
molluscicidal properties (WHO, 1983), which are likely to be cheaper and available on a longer 
term. 

Biological control is a cheap and environmentally attractive alternative to chemical control but 
further work is needed. It may play an important role in the maintenance of control 
programmes but it is important to bear in mind that it may have an effect on non target 
organisms (WHO, 1981). 

Habitat control and appropriate agricultural engineering may be one of the most effective 
methods as its effect is persistent, but it has disadvantages. Removal of water may be effective 
but in many cases can only be considered a temporary measure, particularly as snails can 
survive considerable desiccation (Webbe and Jordan, 1982). Removal of natural habitat is also 
recommended in some instances and clearly this could have considerable bearing on the 
survival of non target species. Other control methods which are important in the longer term 
include curative drugs, and improved water supplies, sanitation and health education. Since 
schistosomiasis considerably slows down economic development in regions where it is endemic, 
it is in fact preferable to prevent its introduction. 

The Mangoky Irrigation Project in south-west Madagascar was one of the six major control 
programmes throughout the world involving the control of the snail hosts. It was set up at the 
same time as the Samangoky Mixed Management Company was developing a vast irrigation 
system to be used for cotton cultivation. The area readily lent itself to the prevention of 
schistosomiasis and to the development of control methods. In 1971 the project was transferred 
to the Madagascan government, a team having been trained to take over the project. At this 
time, the project was considered to have been successful, as snails had been reduced or 
eliminated using the molluscicide Frescon. However, the risk of reintroduction was considered 
to be very high unless certain preventive measures continued to be carried out (Degremont, 
1973). 

-88- 



Fauna 
We are very grateful to D.S. Brown for providing information for this section. 



REFERENCES 

Brown, D.S. (1978). Freshwater mollusks. In: Werger, M.J. A. (Ed.), Biogeography and 

ecology of southern Africa. Part 1 and 2. Monographiae Biologicae 31. Junk, the Hague. 
Brown, D. (1980). Freshwater snails of Africa and their medical importance. Taylor 

and Francis, London. 
Brygoo, E.R. (1965). Les Bilharzioses humaines a Madagascar. Arch. Inst, pasteur 

Madagascar 33: 79-206. 
Brygoo, E.R. (1968a). Les Bilharzioses humaines a Madagascar. In: Sante et Developpement 

I er Congres International des Sciences Medicales de Madagascar. Pp. 1-165 
Brygoo, E.R. (1968b). Temperature and the distribution of human schistosomiasis on 

Madagacar. Bull. Soc. Pathol. Exot. 60(5): 433-441. 
Brygoo, E.R. (1972). Human diseases and their relationship to the environment. In: 

Battistini, R. and Richard-Vindard, G. (Eds), Biogeography and Ecology of 

Madagascar. Monographiae Biologicae 21. Junk, the Hague. 
Brygoo, E.R. and Moreau, J. P. (1966). Bulinus obtusispira (Smith, 1886) hote 

intermediate de la Bilharziose a Schistosoma haematobium dans le nord-ouest de 

Madagascar. Bull. Soc. Path. Exot. 59: 835-839. 
Christie, J.D. (1978). A review of factors determining the spatio-temporal dynamics of 

human schistosoma transmission. Draft report, WHO. 
Degremont, A. A. (1973). Mangoky Project. Campaign against schistosomiasis in the Lower 

Mangoky (Madagascar). Basle: Swiss Tropical Institute. 
Fischer-Piette, E. and Vukadinovic, D. (1973). Sur les mollusques fluviatiles de 

Madagascar. Malacologia 12(2): 339-378. 
Hoffman, D.B., Lehman, J.S., Scott, V.C., Warren, K.S. and Webbe, G. (1979) Control of 

Schistosomiasis. Am. J. Trop. med. Hyg. 28(2): 249-259. 
Perret, P., Egger, M. and Degremont, A. A. (1972). Essai de lutte anti-mollusque par 

augmentation de la biomasse planctonique. Acta Tropica 29: 175-181. 
Pfluger, W. (1977a). Ecological studies in Madagascar of Biomphalaria pfeifferi 

intermediate host of Schistosoma mansoni. 1. Seasonal variations and epidemiological 

features in the endemic area of Ambositra. Arch. Inst. Pasteur Madagascar 45: 79-1 14. 
Pfluger, W. (1977b). Ecological studies in Madagascar of Biomphalaria pfeifferi 

intermediate host of Schistosoma mansoni. 2. Biology and dynamics in the non-endemic 

area of Antananarivo. Arch. Inst. Pasteur Madagascar 46: 241-269. 
Prod'hon, J. Richard, J., Brygoo, E.R. and Daynes, P. (1968). Presence de 

Paramphistomum microbolhrium Fischoeder 1901 a Madagascar. Arch. Inst. Pasteur 

Madagascar 37(1): 27-30. 
Starmuhlner, F. (1962). Voyages d'etudes hydrobiologiques a Madagascar 1958. 

Naturaliste Malgache 13: 53-83. 
Starmuhlner, F. (1969). Die Gastropoden der madagassischen Binnengewasser. 

Malacologia 8: 1-434. 
Starmuhlner, F. (1979). Distribution of freshwater molluscs in mountain streams of 

tropical Indo-Pacific islands (Madagascar, Ceylon, New Caledonia). Malacologia 

18(1-2): 245-256. 
Webbe, G. (1981) The six diseases of WHO. Schistosomiasis: some advances. Brit. Med. 

J. 283: 1104-1106. 
Webbe, G. and Jordan, P. (1982). Control. Chapter 11. In: Jordan, P. and Webbe, G. 

(Eds) Schistosomiasis. William Heineman Medical Books Ltd., London. 
WHO, (1981) Data sheet on the biological control agent, Thiara granifera (Lamarck). 

World Health Organisation, WHO/VBC/8 1.833. 
WHO, (1983). Report of the scientific working group on plant molluscicides. UNDP/World 

Bank/WHO special programme for Research and Training in Tropical Diseases, 

TDR/SCH-SWG(4)/83.3. 
Wright, C.A. (1971). Bulinus on Aldabra and the subfamily Bulininae in the Indian 

Ocean area. Phil. Trans. R. Soc. 260: 299-313. 



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An environmental profile of Madagascar 

V.8. MARINE MOLLUSCS 

There are a number of publications on the marine molluscs of Madagascar (see Brygoo and 
Brygoo, 1978; Mars et al., 1972). About 1020 species have been described. The fauna is 
generally considered to be typical of the Indo-Pacific although the possibility of a separate 
classification has been raised (Mars et al., 1972). However, consideration of the comparatively 
extensive studies which have been made on the genus Strombus and the family Cypraeidae 
suggests that there is little basis for this. 

Brygoo and Brygoo (1978) provide a checklist of 66 Conus species and 56 Cypraea species. 
Cowries are abundant around Nosy Be and include species such as Cypraea nucleus, C. 
diluculum. C. onyx. C. oweni. C. lamarcki, and C. chinensis (Magnier, 1981). At least 15 
families of opisthobranchs are represented in Madagascan waters, including 9 tectibranch 
families, 6 nudibranch families and one family of Opisthopneumones (Vicente, 1966). The 
only marine mollusc which appears to be endemic to the region (Madagascar to Seychelles) is 
Delessert's Volute, Lyria (= Voluta) delessertiana, a small red-orange species, which is said to 
be found off Dzamandzar (Magnier, 1981). 

Ecological studies have been carried out only recently. These are outlined in Mars et al. 
(1972), where it is pointed out that some rare genera have recently been found in the region of 
Toliara: Berthelinia in the Caulerpa zone, and the commensal gastropod Caledoniella 
montrouzieri which parasitises the abdomen of stomatopod crustaceans. 

Mars et al. (1972) point out the importance of reference collections for use in future research 
work. An 'accord' has been drawn up stating that specimens would be sent to the Museum 
National d'Histoire Naturelle at Paris. Following systematic revision of each family, three 
reference collections would be created, one for the museum, one for the Station marine 
d'Endoume at Marseilles and one for the marine laboratory at Toliara. It is not known to what 
extent this has been carried out. 



Food Species 

Decary (1950) reported that in spite of their variety and abundance, marine molluscs were not 
used very much for food except in certain areas. However Rabesandratana (1985) noted that 
with the increasing difficulty of obtaining meat, molluscs had become more important as food 
items; Table 8, taken from Rabesandratana (1984 and 1985), lists over thirty mollusc taxa 
which are exploited as a food source in Madagascar. In the north-west, the Sakalava people 
occasionally eat Area species (Kodiva) while between Antseranana and Toamasina large Nerita 
(Sifotro mamy) are collected at low tide when they come out of the sand. Tridacna 
(Hima), Turbo (Betampy) and Pleuroploca species are also eaten. Rabesandratana (1972) gives 
further examples on Nosy Borah. In the 1960s it was common to see children at low tide 
collecting a variety of molluscs (Conus, Strombus. Area, Cardium, Venus. Tridacna) which are 
an important part of the local diet, often replacing meat. In the extreme south, the Antandroy 
people on the coast collect Cellana capensis; around Toliara the Vezo fishermen 
collect Pleuroploca, Ceriihium, and Murex species which they eat cooked. In the region of 
Lamboharanana (baie des Assassins) people collect chitons for food. Mussels may not be 
collected less than 3.5 cm in length (Randrianarijaona and Razafimbelo, 1983). 

a. Oysters 

Oysters (Papakiny, Saja) are certainly the most important food mollusc both locally, e.g. on the 
coast at Androy and in the region of Taolanaro, or sent inland to the large towns. 
Rabesandratana (1972) provides a bibliography of species descriptions for this group. The two 
most important edible species are: 

Crassostrea cucullata (Born), the most abundant species and found on almost all coasts; has a 
wide Indo-pacific range and is cultured in many countries; varies in form according to 
habitat. Ostrea vitrefacta (Sowerby), abundant in the region of Mahajanga, may be a variety of 
this species. 

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Fauna 

Crassostrea margaritacea (Lamarck), the 'huitre pied de cheval de Madagascar'; found also in 
South Africa; occurs mainly around Taolanaro. 

Natural oysterbeds in the south and south-west of Madagascar have been studied for several 
years. On the south-west coast, the beds of C. cucullata between Morombe and 
Lamboharanana which are exploited commercially have practically disappeared, and have been 
completely exhausted in the region between Ifaty and the Manombo estuary to the north of 
Toliara. South of Toliara, in the region of Sarodrano, there is an important bed which is 
partially protected by a provincial decree for the purposes of research and aquaculture 
experiments. Provincial decree no. 054- AG of 6 May 1966 prohibits collection of oysters 
throughout the year in the bay of Sarodrano and during the breeding season in the area around 
Faritany. Studies are being carried out and it has been recommended that the results should be 
made known widely (Randrianarijaona and Razafimbelo, 1983). The decree of 5 June 1922 
prohibits collections of oysters with a diameter of less than 4 cm. 

To the south of the bay of Saint-Augustin, oyster population density is much higher. The only 
other important beds are in the region of Androy: Lavanono, Fanambosa and Ankatravitra. By 
the 1970s the bed at Ankatravitra had already started to decline. 

Beds of C. margaritacea are found only between Evatraha and Ambinanibe at Taolanaro. This 
species is usually collected by skin divers, and although exploitation is not as intense as if other 
methods were used, collectors have commented that oysters are becoming rarer. C. cucullata is 
usually collected with hammers and chisels. Often whole chunks of substrate are taken and the 
oysters are separated off later. In general, the methods used lead to great habitat destruction 
and prevent the settlement of juveniles, since the substrate becomes silted up and overgrown 
with diatoms. Such methods could result in beds disappearing within only a few years. Major 
action needs to be taken. Recommendations were made in 1972 but it is not known if these 
have been implemented: 

1. A closed season should be imposed from November to the end of March, during the 
breeding season. 

2. The size limit which was imposed under a decree in 1922 should be strictly adhered to: 
4 cm longitudinal axis, a size generally reached at 3 years. 

3. Aquaculture must be encouraged; trials were being carried out with C. cucullata in the early 
1970s. 



Other uses 

Decary (1950) describes many other uses for marine shells. Near Maroantsetra and Mananara, 
the large valves of Pinna shells ('lelabasy') are used for bailing out canoes and as plates. In the 
1920s Pinna nigra was collected from sea grass beds for its pearls and byssus threads which 
were used for making silk. Shells are used for weighting fishing nets. Many molluscs are used 
as fish bait, particularly Cerithium palustre. Shells are used to decorate houses, and in 
Mahajanga province jewellery is made from small Trochus and Gibbula shells. The 'felana 1 or 
'fela' is a white disc made from the spire of a Conus shell which was worn on the head by 
warriors in the tribes of the west and north; it is now mainly worn for decoration by women, 
particularly professional dancers known as Sahafatra in Farafanga province. Small cowries, 
('androvo', 'horohoro') are used in games. Cypraea annulus is considered a talisman; in the past 
it was exported to North Africa for use as currency. In the south-west, the operculums 
of Pleuroploca (= Fasciolaria) and Mure x ('fimpy') are used by the Mahafaly and Sakalava 
people for making a perfume used in sorcery, and shells are used in other magical rituals by 
the Sakalava people. 

The conch, Charonia tritonis ('antsiva', Mavabody') was used as a trumpet in war and on other 
occasions, for example to call the wind if becalmed at sea, at burials, as a warning that locusts 
were spreading or a storm was coming. Pleuroploca and Cassis ('antsiva boriborivody') shells 
were also occasionally used as trumpets. Shell trumpets are still used sometimes. At Androy 
there are two forms of trumpet: Charonia ('antsiva lahy') is the male, is large and gives a 
powerful sound; Pleuroploca ('antsiva vavy') is the female, smaller and quieter. The male 

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An environmental profile of Madagascar 

trumpet can be used on its own or with the female; the female can only be used with the male 
and never on its own. Charonia tritonis is used as a trumpet in different countries throughout 
the Indo-Pacific. It is also a very popular species in the ornamental shell trade. It is listed as 
Rare in the IUCN Invertebrate Red Data Book (Wells et al., 1983) as its population density is 
naturally low, and there are fears that it could be threatened by intensive exploitation. 



Exports 

Madagascar exports substantial quantities of marine shells and shell products, the most 
important of which is mother-of-pearl from the species Turbo marmoratus and Trochus 
niloticus ('betampy'). These species are most abundant on the west coast (both are reef 
dwellers) and are collected for export around Toliara. Export figures for the years 1976-84 are 
included in Table 10. Exports principally go to Europe (France, Italy, Spain, United Kingdom) 
and occasionally to India, Kenya and Japan (Rabesandratana, 1985). Before the second world 
war, 150-200 tonnes were collected annually with evidence of overexploitation of large 
specimens (Decary, 1950). In several parts of the world these species have been overfished and 
legislation has been introduced to control the fishery (Wells, 1980). The four important 
mother-of-pearl species, Trochus niloticus, Turbo marmoratus, Pinctada margaritifera and P. 
maxima, are categorised as Commercially Threatened. The current status of the fishery in 
Madagascar is not known, although reported exports have declined considerably from 
1978/1979 to 1983. 

Second in importance to mother-of-pearl is the Bullmouth Helmet Cypraecassis rufa which is 
principally exploited along the south-west coast from Morombe to Anakao, south of Toliara, 
being rare further north. Two forms are collected, one, the 'male' shell, is exported, 
principally to Italy, for the production of shell cameos, the second, the 'female' shell, is used 
for making ash-trays. Export figures for the years 1976-84 are given in Table 10. 
Rabesandratana (1985) quotes mean weight estimates for a 'male' shell of 0.8 kg, for a 'female' 
shell of ca 0.3 kg. From these, the numbers exported from Toliara in the years 1979-82 would 
be likely to lie between 77 000 (if all 'males') and 185 000 (if all 'females'). 

Considerable numbers of other species, principally gastropods, are also exported; a breakdown 
of the figures for 1978 and 1979 is included in Table 11; Rabesandratana (1985) considers that 
several species are probably being over-collected and recommends some form of control. 

Decary (1950) reports that pearls are occasionally found in Pinctada (Meleagrina) 
margaritifera, M. occa and M. irradians but since the Madagascans do not have a tradition of 
diving for pearls only the easily accessible specimens are taken. Sometimes pearls are exported 
to Sri Lanka. The decree of 23 August 1929 imposes controls on collection of pearl oysters, 
mother-of-pearl shells and sponges (Randrianarijaona and Razafimbelo, 1983). 

a. Pearl oysters {'Meleagrina' occa, M. irradians): 6 cm minimum size, measured from inside 
edge of valves across largest diameter. 

b. Mother-of-pearl oyster (M. margaritifera): 10 cm minimum size. 

c. Turbos or Bourgos (Turbo marmoratus): 55 mm minimum size; 140 mm maximum size. 

d. Trochus or Trochas (Trochus niloticus): 1 10 mm minimum size. 

The substrates of pearl oysters (corals, etc.) must be thrown back into the sea. 

The IUCN Invertebrate Red Data Book (Wells et al., 1982) points out that although most 
marine molluscs are unlikely to be threatened with extinction through human activities, local 
overexploitation can easily occur. This is particularly evident with the giant clams in the 
family Tridacnidae. It is recommended that commercial exploitation of any marine mollusc is 
carried out under an appropriate management strategy. Further information is required on the 
shell trade, and it might be necessary to control collection of Lyria delessertiana if this proves 
to be endemic. 



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Fauna 

TABLE 8. MOLLUSCS EXPLOITED FOR FOOD IN MADAGASCAR 

North- South- North- 
west west east 
GASTROPODA 

Patellidae 

Cellana capensis ++(*) 

Turbinidae 

Turbo marmoratus +++ +++ +++ 

Turbo coronatus ++ ++ ++ 

Turbo argyrostomus ++ ++ ++ 

Turbo imperialis +++ ++ 

Naticidae 

(species not given) ++ ++ 

Neritidae 

Nerita albicilla ++ ++ 

Nerita undata ++ ++ 

Littorinidae 

Littorina kraussi ++ 

Littorina acabra ++ 

Conidae 

Conus lividus ++ +++ 

Conus tessulatus ++ +++ 

Fasciolariidae 

Pleuroploca trapezium +++ +++ +++ 

Pleuroploca filamentosa + ++ 

Muricidae 

Murex ramosus +++ +++ +++ 

Drupa morum ++ 

Stombidae 

Strombus gibberulus +++ 

Cerithiidae 

Pyrazus palustris 

BIVALVES 

Arcidae 

Anadara natalensis 
Cardiidae 

Cardium sp./spp. +++ 

Mytilidae 

Septifer bilocularis ++ 

Modiolus sp./spp. +++ 

Ostreidae 

Crassostrea cucullata +++ +++ +++ 

Crassostrea margaritacea +++(#) 

Pteriidae 

Pine tad a margaritifera 
Tridacnidae 

Tridacna squamosa 
Veneridae 

Gafrarium pectinatum 

CEPHALOPODA 

Octopus sp./spp. 

Loligo sp./spp. +++ +++ +++ 

Sepia sp./spp. +++ + ++ 

+++ much sought after; ++ harvested; + harvested occasionally. 
* extreme south; # south-east (Taolanaro). 
Taken from Rabesandratana (1984). 

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An environmental profile of Madagascar 

TABLE 9. EXPORTS OF SHELLS, 1976-84 (TONNES) 

1976 1977 1978 1979 1980 1981 1982 1983 1984 



Turbo marmoratus 
















Toliara 3 


- 


89.3 


46 


10 


37 


14 


5# 


Madagascar 13 87.6 


53 96.2 


94.2 


39.2 


15 


42.4 


14* 




Cypraecassis rufa 
















Toliara 3 


- 


14.55 


23.08 


10 


14 


- 


3# 


Total shells (including C. 


rufa, excluding T. 


marmoratus) 










Toliara 3 


- 


46.2 


48.3 


78.3 


153.3 


125.6 


11.7* 


Madagascar* 3 61.4 


82.4 63.9 


37.7 


103.3 


71.1 


63.9 


10.6* 





# = First half of year; * = figures for five months only; - = no information; 

a = source Bureau de Recette des Douanes de Toliara; Service Provincial de Commerce de 

Toliara 
b = source Institut National de la Statistique et de la Recherche Economique (I.N.S.R.E.) 



TABLE 10. EXPORTS OF SHELLS FROM TOLIARA, 1979 AND 1980 (KG) 

Species 1979 1980 

Cypraea spp. 
Cypraea caputserpentis 
Cypraea histrio 
Cypraea scurra 
Cypraea tigris 
Conus spp. 
Charonia tritonis 
Lambis lambis 
Lambis truncata 
Oliva spp. 
Terebra spp. 
Polynices spp. 
Vasum ceramicum 
Cypraeacassis rufa 
Murex ramosus 
Turbo bruneus 
Turbo marmoratus 
Bursa lampas 
Bursa bubo 
Strombus lentiginosus 
Clanculus pharaonius 
Pleuroploca trapezium 
Atrina vexillum 
Macrocallista nimbosa 
Tridacna elongata/gigas 

Source Bureau de Recette des Douanes de Toliara; Service Provincial de Commerce 
de Toliara 

Taken from Rabesandratana (1985). 



1000 


2950 


125 


- 


130 


110 


622 




480 


200 


412 


100 


169 


101 


791 


448 


558 


766 


300 




2040 


1000 


570 


500 


100 




750 


2100 


8660 


13087 


520 


200 


1050 


2600 


252 




591 


295 




250 


20 


- 


600 


500 


76 


215 


100 




710 


500 



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Fauna 
REFERENCES 

Brygoo, J. and Brygoo, E.P. (1978). Cones et porcelaines de Madagascar. Arch, de 

I'institut Pasteur de Madagascar Numero Special. 
Decary, R. (1950). La faune malgache, son role dans les croyances et les usages 

indigenes. Payot, Paris. 236 pp. 
Magnier, Y. (1981). Merveilles sous-marines. Chapter 7. In: Oberle, P. (Ed.). 

Madagascar: un sanctuaire de la nature. Lechevalier, Paris. 
Mars, P., Salvat, B. and Thomassin, B. (1972). La faune malacologique littorale marine de 

Madagascar. In: Comptes rendus de la Conference Internationale sur la conservation de la 

nature et de ses ressources a Madagascar. Tananarive. 7-11 October. IUCN Publ. N.S. 

Suppl. Doc. 36. 
Rabesandratana, H.D. (1972). Evolution de quelques gisements d'huitres dans le sud et le 

sud-ouest de Madagascar. In: Comptes rendus de la Conference internationale sur la 

conservation de la nature et de ses ressources a Madagascar. Tananarive, 7-11 October. 

IUCN Publ. N.S. Suppl. Doc. 36. 
Rabesandratana, H.D. (1984). Letter to C. Sheppard / IUCN with enclosures including 

documentation on submission of proposals to establisha Marine Park at Tulear (Toliara) 

and data on collection of molluscs in Madagascar. 
Rabesandratana, H.D. (1985). About some reef utilisations in Madagascar. Proc. 5th Int. 

Coral Reef Cong., Tahiti 6: 661-668. 
Randrianarijaona, P. and Razafimbelo, E. (1983). Rapport national pour Madagascar. 

Report prepared for UNEP Regional Seas East Africa programme. 
Vicente, N. (1966). Contribution a l'etude des Gasteropodes opisthobranches de la region 

de Tulear. Ann. Univ. Mad. 4: 97-142. 
Wells, S.M. (1980). The International Trade in Marine Shells. Report to TRAFFIC 

International (Conservation Monitoring Centre, Cambridge). 
Wells, S.M. (1981). International Trade in Ornamental Shells. IUCN Conservation 

Monitoring Centre, Cambridge. 
Wells, S.M., Pyle, R.M., Collins, N.M. (1983). The IUCN Invertebrate Red Data Book. IUCN 

Conservation Monitoring Centre, Cambridge. 






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An environmental profile of Madagascar 

V.9. NON-MARINE CRUSTACEANS 

The non-marine Crustacea of Madagascar remains little studied, although there are known to be 
many endemic species. Details of some of the more interesting groups are given in Appendix 
2. Several species are restricted to phreatic (groundwater) habitats including copepods 
(Cyclopidae, Harpactidae), ostracods and syncarids. Since rough substrates are most favourable 
for these species, they tend to be found mainly in the fine gravel of mountain torrents 
(Paulian, 1961). 

Fisheries 

Freshwater Crustacea in the order Decapoda are often harvested for food; the most important 
families taken are the Palaemonidae (freshwater shrimps) and the Parastacidae (freshwater 
crayfish), although freshwater crabs (family Potamonidae) and shrimps of the family Atyidae 
are also taken. 

a. Family Palaemonidae 

The larger freshwater shrimps are a very popular food and are known locally as 'patsa', 
'makamba' and 'tsivakiny' (Decary, 1950; Griveaud, 1981) (prawns are 'tsitsika' (Kiener, 
1963)); some of the Macrobrachium prawns are known as 'orana' or 'camaron' such 
as Macrobrachium australe, M. idae (said to be much appreciated by Europeans; 
with Metapenaeus monoceros and Palaemon concinnus it dominates the fishery in the 
Pangalanes), and M. lepidactylus (Madagascar scale prawn, Bouquet malgache, also referred to 
as Palaemon hilgendorfi) (Louvel, 1930; Moulherat and Vincke, 1968). In the past, freshwater 
shrimps were brought as gifts to women in confinement; nowadays visitors bring a small sum 
of money called 'vola amidy patsa' or 'money to buy shrimps' (Decary, 1950). Shrimp are 
caught in the lakes and marshes of the west and in the Pangalanes-Est. Prawns are mainly 
found in rocky parts of rivers at medium and low altitudes, and are caught by women using 
molluscs as bait. A particularly large species (scientific name not identified) found in forest 
streams with long claws and bright blue, green and pink in colour is called 'rangaza', 'rakaho' 
or 'rafitrako' (Kiener, 1963). 

b. Family Parastacidae 

Astacoides madagascariensis is the only representative of the family Parastacidae in Africa, 
and is in a monotypic genus endemic to Madagascar, apparently most closely related to 
Australian and South American forms (Griveaud, 1981). It is absent from the west of the 
country (Decary, 1950). All subspecies of Astacoides are of a size that makes them valuable 
food items. In the 1950s, they were collected quite intensively to the east of Fianarantsoa and 
around Antananarivo but already seemed to be becoming rarer. They were either brought to 
the town alive for sale in the markets or cooked and sold to travellers on the railway stations on 
the line from Antananarivo to Toamasina. A variety of collecting methods are used (Decary, 
1950). They are considered to be declining due to progressive deforestation, although at the 
beginning of the 1960s some areas, especially the rivers flowing down the easten scarp of the 
island, e.g. Mandraka (Ambatolaona) and Ankaratra (Ambatolampy), had large populations 
(Kiener, 1963). Known as 'oram-bokoka' or 'orambato' (Kiener, 1963), they are forest stream 
dwellers and would probably not lend themselves to artificial culture (Hobbs, 1983). However, 
experiments on crayfish culture using a variety of species suggest that it might be worth 
attempting to raise them artificially (Brinck, 1983). 



Threats 

The IUCN Invertebrate Red Data Book (Wells et al., 1983) discusses the types of human 
activities which may threaten freshwater crustaceans, and gives several examples. Habitat 
alteration and pollution are probably the major threats. The species which may be most 
vulnerable in Madagascar are those of interstitial, subterranean and brackish water habitats 
(Dussart, 1983). Interstitial and subterranean forms may be threatened by pollution. There are 

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Fauna 

a number of interesting endemic taxa (e.g. Anopsilana, Caridina troglophila, Parisia edentata, 
P. macrophthalma, P. microphthalmia and Typhlopatsa pauliani) restricted to small, and 
therefore particularly vulnerable, caves (Paulian 1983). Moss and epiphytic lichens also seems 
to be important habitats for crustaceans, presumably because they provide a high level of 
humidity during the day (Paulian, 1961). Burrowing and cave crustaceans are threatened in 
many parts of the world, and examples are given in the IUCN Invertebrate Red Data Book. 

In many countries, native crayfish are coming under increasing threat through a combination 
of factors including overexploitation, pollution and competition with introduced species. Since 
there is evidence that the native Madagascan species are a valuable resource, efforts should be 
made to ensure that stocks are properly managed. Fishing pressure on the native species is 
already heavy, and certainly destructive. Regulations should be introduced to limit catches and 
to protect the reproductive stages, possibly through closed seasons and minimum sizes (Paulian, 
1983b). 

We are very grateful to Dr P. Brinck, Dr B. Dussart, Dr D.G. Frey, Dr H.H. Hobbs, Dr 
L. Holthuis, Dr P. Lake and Dr R. Paulian for assisting with this section. 



REFERENCES 

Barnard, K.H. (1958). Terrestrial amphipods and isopods from Madagascar. Mem. Inst. 

Sci. Mad. A (12): 67-1 11. 
Barnard, K.H. (1960). A new species of Calmanesia (Isopoda Terrestria) from 

Madagascar. Mem. Inst. Sci. Mad. A (14): 59. 
Bott, R. (1965). Die Siisswasserkrabben von Madagaskar (Crustacea, Decapoda). Bull. Mus. 

Natn. Hist. Nat., Paris Ser. 2. 37(2): 335-350. 
Brinck, P. (1983). In litt. 
Coutiere, (1901). Les Palaemonidae des eaux douces de Madagascar. Ann. Sci. Nat. 

Zool. (8)12: . 
Decary, R. (1950). La faune malgache, son role dans les croyances et les usages 

indigenes. Payot, Paris. 236 pp. 
Dussart, B.H. (1982). Crustaces copepodes des eaux interieures. Faune de Madagascar 

58: 146 pp. 
Dussart, B.H. (1983). In litt. 24 March. 
Germain, L. (1935). Etude de la reserve naturelle du lac Manampetsa. Ann. Sci. Nat. Zool. 

18:421-481. 
Griveaud, P. (1981). Les invertebres: un univers prodigieux. Chap. 3. In: Oberle, P. 

(Ed.), Madagascar: un sanctuaire de la nature. Lechevalier, Paris. 
Hobbs, H.H. (1983). In litt. 3 May. 
Holthuis, L.B. (1964). The genus Astacoides Guerin (Decapoda Macrura). Crustaceana 

6(4): 309-318. 
Holthuis, L.B. (1965). The Atyidae of Madagascar. Mem. Mus. Natn. Hist. Nat. Serie A 

33(1): 1-48. 
Kiener, A. (1963). Poissons, Peche et Pisciculture a Madagascar. Centre Technique 

Forestier Tropical, Nogent-sur-Marne, France. 
Legendre, R. (1972). Apercu sur quelques formes relictes et interessantes des 

invertebres terrestres et d'eaux douces de madagascar (mollusques exceptes). Comptes 

rendus de la Conference Internationale sur la conservation de la nature et de ses ressources 

a Madagascar, Tananarive. 7-11 Octobre. IUCN Publ. N.S. Suppl. Doc. 36. 
Louvel, M. (1930). Sexploitation des eaux douces de Madagascar (peche et 

pisciculture). Tananarive, Gouvernement general de Madagascar et dependances, 52 pp. 
Monod, T. (1935). Crustacea. In: Germain, L. (1935). Etude de la reserve naturelle du lac 

Manampetsa. Ann. Sci. Nat. Zool. 18: 421-481. 
Moulherat, J.L. and Vincke, M. (1968). Etude en vue du developpement de la peche au 

Pangalanes-Est (zone Tamatave-Andevoranto). Madagascar, Centre Technique Forestier 

Tropical, 195 pp. 
Paulian, R. (1961). La zoogeographie de Madagascar et des iles voisines. Faune de 

Madagascar 13: 1-485. 
Paulian, R. (1983a). In litt. 22 April. 

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An environmental profile of Madagascar 

Paulian, R. (1983b). In lilt. 28 July. 

Paulian, R. and Delamare Deboutteville, C. (1956). Un Cirolanide cavernicole a 

Madagascar. Mem. Inst. Sci. Mad. Ser. A 11: 85-88. 
Paulian de Felice, L. (1950). Oniscoides nouveaux de Madagascar. Naturaliste malgache 

2: 101-106. 
Roux, J. (1934). Macroures d'eau douce de Madagascar et des lies voisines. Faune des 

colonies francaises 5(8): 529-547. 
Ruffo, S. (1958). Amphipodes terrestres et des eaux continentales de Madagascar, des 

Comores et de la Reunion (Etudes sur les Crustaces, Amphipodes -L.). Mem. Inst. Sci. 

Mad.: 35-66. 
Ruffo, S. (1979). Descrizione di due nuovi anfipodi anoftalmi delF Iran e de Madagascar 

(Phreamelita paceae n. gen., n. sp., Dussartiella madegassa n. gen., n. sp.). Boll. Mus. Civ. 

Stor. Nat. Verona 6: 419-440. 



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Fauna 

V.10. MARINE CRUSTACEANS 

Like most other Madagascan marine invertebrates, the marine crustaceans are mainly typical of 
the Indo-Pacific fauna; some groups have been fairly well studied but there is still much work 
to be done. Many of the publications up to 1974 are listed in ORSTOM (1973) and ORSTOM 
(1975). It has not been possible to review all the studies that have been carried out on marine 
crustaceans but a few examples are given below, and the main commercial crustacean fisheries 
are discussed. 

Seventy-three species of crabs in the family Portunidae have been described (Crosnier, 1962), 
collected in the course of studies on peneids. Collecting was carried out on all coasts except 
the east; 23 of the species found occur in South African waters, 33 in Australasian waters, 18 
in Hawaii, and 33 in Japan. Five species new to science were described. The study confirmed 
the fact that the marine fauna of the north-west and west coasts of Madagascar and the 
Comoros is very rich in species and homogeneous; in the south and south-east, the waters art 
colder and there is a rather different and less diverse fauna. 

Forty-one species of crab in the family Grapsidae and 19 in the family Ocypodidae (which 
includes the ghost crabs Ocypode and the fiddler crabs Uca) have been described (Crosnier, 
1965), but these lists are almost certainly incomplete, particularly for the smaller forms. Six 
new species were described. All except three species were intertidal. This study illustrated 
that most Madagascan marine species fall into three groups: 

1. Species with Indo-Pacific distributions 

2. Species with distributions extending as far as South-East Asia or India. 

3. Species limited to Madagascar and the East African region. 

Species which are apparently endemic may only be so through lack of collection in other 
places. Rare exceptions concern some of the species from the region of Taolanaro and St. 
Lucia in the south-east; it is possible that some of these, such as Ocypode madagascariensis, 
are true endemics (Crosnier, 1983). Studies on the families Xanthidae, Trapeziidae, Carpiliidae 
and Menippidae have been carried out and will shortly be published; more than 200 species 
have been recorded (Serene, in press). There are also a number of endemic marine phreatic 
species, revealed by samples taken from the lagoon across the 'cordon littoral' at Maroantsetra. 
These may have extremely restricted distributions if they are comparable to endemic species 
found in similar habitats on Reunion (Paulian, 1961). 

The Coconut crab Birgus latro has been recorded from the Comoros (Fourmanoir 1953) and 
may possibly occur on some of the smaller islands off the Madagascar coast. It is listed in the 
IUCN Invertebrate Red Data Book as Rare, and there is evidence that it is declining in some 
parts of its range. Information on its presence in Madagascar is required. 



Lobster fishery 

Three reports on the lobster fishery are available (Charbonnier and Crosnier, 1961; Pichon 
1964; Fourmanoir el al., 1960). In the east, north-west and west, where the coast is mainly 
coralline, the following species predominate: Panulirus versicolor (Latr.) Painted Crayfish 
(Indo-Pacific); P. omatus (Fabr.), the largest species, weighing up to 3 kg.); P. japonicus 
(Siebold) Japanese Spiny Lobster, Langouste Japonaise; in addition P. penicillatus was found to 
be rare on Nosy Be (Pichon, 1964). 

These species are usually found at depths of less than 4 m. Their reef habitat precludes the use 
of nets and traps and they are taken only by divers and spear fishermen, although this is said to 
be illegal (see below). There is no substantial fishery. 

On the rocky coasts of the south and south-east, Panilurus hommarus (L.) (= P. burgeri) and P. 
penicillatus are found (Charbonnier and Crosnier, 1961). 



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An environmental profile of Madagascar 

These two species are caught easily and an artisanal fishery has developed along the coast 
between Cap Ranavalona and Manantenina. P. penicillatus is quite rare and constitutes more 
than 1% of the fishery. The principal fishing centres are the bay of Itaperina and Sainte Luce. 
Lobsters are caught using traps placed either by wading out or using canoes made from tree 
trunks. Most fishing is carried out at depths of less than 8 m; attempts at depths of 25-50 m 
were unsuccessful (Fourmanoir et al., 1960). Mussels are the most suitable bait, but in the 
more populated areas these have been depleted, and limpets {Patella sp.) and acorn barnacles 
(Balanus sp.) are used instead. Mussels are still abundant in areas offshore but are inaccessible 
here because of the sea swell. The number of fishing days per year is very small on account of 
winds, heavy swell and the instability of the canoes; in general, only sheltered sites are 
exploited. In 1958, 24.5 tonnes were produced; in 1959, 21.5 tonnes. At the beginning of the 
1960s the 'Division des Peches maritimes du Service de l'elevage et l'lnstitut de recherches 
scientifiques a Madagascar' carried out a survey of the Taolanoro and Sainte Luce areas to see 
if an extension of the fishing zone would be feasible. However it was found that the range of 
the lobsters is restricted to a very narrow coastal band. An increase in harvest could have been 
achieved by using motor boats to get to the more inaccessible sites, but in the 1960s the local 
markets in Madagascar were already saturated. A freezer plant would have to be installed if an 
export trade were to develop; this would necessitate a minimum production of 10 tonnes per 
month, which would be difficult to achieve. Furthermore, an increase in fishing intensity 
would lead to even greater depletion of the bait species. It was therefore concluded that 
further intensification of the lobster fishery would not be a viable proposition (Charbonnier 
and Crosnier, 1961). Attempts were made to improve yields using artificial shelters, but these 
were not successful (Pichon, 1964). Frozen tails and living lobsters have been exported to 
France on occasions, but quantities are not available and it is not known if exports continue. 
The decree of 27 December 1962 prohibits the fishing, sale, transport and processing of 
lobsters from 1-30 April each year (Randrianarijaona and Razafimbelo, 1983). Throughout the 
year it is forbidden to sell, transport or sell ovaried female lobsters and any lobsters less than 
20 cm in length (measured from the end of the telson to the end of the rostrum excluding the 
antennae). The decree of January 1921 prohibits the use of all collecting methods apart from 
traps and nets. These methods permit the release of ovaried females, unlike harpooning. 
There is no comment on the fact that harpooning is the method used in the east, north-west 
and west. Studies on deep-sea Palinurus have been carried out by Plante and Moal. 



Crab fishery 

Crabs are known locally as 'foza'. Scylla serrata, the mud crab (crabe de mangrove), which 
has a range throughout the Indo-Pacific, has long been fished in Madagascar but a commercial 
fishery for export has only recently developed, mainly as a sideline by the shrimp companies. 
The fishery at Antseranana was studied in 1976 by Le Reste, Feno and Rameloson. Adults and 
very young juveniles and larvae live in shallow open water, juveniles are found in brackish 
estuaries and subadults in the mangroves. These habitats are found primarily on the west coast 
(98% of Madagascan mangroves are found here) and crab fisheries have developed at Toliara, 
Mahajanga and Antseranana. Young crabs move into estuarine waters at a size of 2-8 cm 
(cephalothorax width). On attaining a size of 10 cm they move to the mangroves where mating 
occurs, and then into the sea where the larvae are released. There appear to be two breeding 
seasons, one in the dry season and one in the wet season. Crabs feed on shrimps, crabs, fish 
and bivalves, and although occasionally found on mud, they usually inhabit burrows in the 
mangroves. 

Crabs are caught while they are in the mangroves, probably aged about 5 months; since the 
females move into the mangroves later than the males, a higher percentage of male crabs are 
caught. They are caught in their burrows, a method known as 'peche au trou' which has been 
shown to be most efficient. In some areas, however, they are reported to be caught in sea grass 
beds. At Antseranana they are caught mainly in the warm and wet season (i.e. October to 
April, with a peak in December/January) when they are most abundant, unless rainfall is 
sparse when they are more abundant in the dry season and caught then. At Toliara the 
maximum catch is taken in April; at Mahajanga the fishing season runs from March to July. 
The significance of these differences is not known. At Antseranana the crabs show two peaks 
of abundance, one in the wet season and one in the dry, which are considered to be linked to 

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Fauna 

the two peaks in primary production, typical of tropical waters. Prior to export, the crabs are 
kept in mud but studies have shown the mortality to be fairly high. Males give the best yield 
of flesh and are particularly valuable on account of their large claws which are considered a 
delicacy. Crabs for export are cooked and their shells removed. 

Total production in the early 1970s was as follows: 

1970 182 tonnes 

1971 258 " 

1972 161 " 

1973 111 " 

Exports have probably ceased altogether now. The decline between 1971 and 1973 was 
considered to be due to a decrease in fishing effort rather than overexploitation, as crabs are 
relatively expensive and both foreign and local markets had declined. FAO statistics do not 
record Madagascar as a producer for the years 1977-1980. The 1976 survey recommended that 
studies should be carried out on the Toliara and Mahajanga fisheries and suggested that culture 
of this species might be possible but would require considerable further research, and would 
probably present many problems (see below). It was concluded that the traditional fishing 
methods used would be difficult to better, that there was little likelihood of overexploitation 
since the crabs grow fast and are only exploited in the mangroves where there are a higher 
percentage of males, and that the large males are the most profitable specimens to take. 

In other countries, the mangrove crab is cultured as a subsidiary crop to milkfish in 
brackish ponds. The ponds are seeded with small crabs which reach marketable size within 
six months (Warner, 1977; Bardach et al., 1972). 



Shrimp fishery 

This is the largest crustacean fishery, and is of major economic importance; shrimp are the 
fifth most important export commodity. Most exports go to Japan and the USA which are 
the main shrimp consumers. The fishery began commercially in 1967 off the north-west 
coast and developed rapidly until there were more than 40 vessels involved in 1975 
(Marcille 1978). Numerous publications are available on this subject as the fishery has been 
well studied by ORSTOM; some of these are listed in the bibliography. 

The following species are the most important (details from Holthuis (1980) and further 
details on these species are given in Marcille (1978)): 

Penaeus indicus Milne Edwards 1837. The Indian White Prawn, Crevette Royale Blanche, 
'makamba' (Sakalava language), 'patsa' (Hova language). Occurs thoughout Indo-West 
Pacific. Makes up 67% of the total shrimp catch in Madagascar, equivalent in 1974 to 3000 
tonnes. 

P. semisulcatus De Haan, 1844. The Green Tiger Prawn, Crevette Tigree Verte. Indo-West 
Pacific, and has reached eastern Mediterranean through the Suez canal. 11% of Madagascar 
shrimp catch. 

P. monodon Fabricius, 1798. The Giant Tiger Prawn, Crevette Giante Tigree. Indo-West 
Pacific. 3% of shrimp catch. 

Metapenaeus monoceros (Fabricius, 1798). The Speckled Shrimp, Crevette Mouchetee, 
'patsanorana' (name used also for other species). Indo-West Pacific and eastern 
Mediterranean. 19% of catch. 

P. japonicus (Bate) is fished in very small quantities. 

Fishing is coastal and carried out at depths of 5-25 m in sandy and muddy areas. The 
continental shelf of Madagascar is poorly developed except in the region of Taolanaro and 

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An environmental profile of Madagascar 

on the west coast where the fisheries are concentrated. The commercial fisheries are 
concentrated between the Mangoky estuary and cap Saint Sebastien (see map in Marcille, 
1978). Two kinds of fishery have developed: 

An artisanal fishery consisting of shore barriers, which catches mainly immature 
shrimps in estuaries near mangroves in the intertidal zone. 

An industrial fishery operating with a trawler fleet and freezer plant which catches 
adult shrimps in depths of 3-10 m. The products are processed in four factories. 
Research has been carried out on the production of edible products using waste from 
the shrimp industry (Frontier Albon, 1972). 

The ORSTOM centre at Nosy Be has collected statistics since the beginning of the fishery. 
This has permitted a study of the evolution of stocks over this period, during which there 
was considerable intensification of the fishery. The results are given in Marcille (1978), 
where models have been produced which could be used to help determine the maximum 
fishing effort that shrimp stocks could support. The study concludes that some form of 
fishery management is required, since in 1974 the catch declined despite increased fishing 
effort (in 1975 catches increased slightly - see figures in Marcille, 1978). The following 
recommendations were made: 

For the artisanal fishery, the size of the gaps between the lattices in the shore barriers 
should be increased from 7.5 mm to 1 1 mm to increase selectivity. 

For the industrial fishery, a closed season should be introduced throughout the 
fishing area for a short period (1-2 months) in January/February in order to protect 
the young. A closed season was implemented on part of the north-west coast near 
Mahajanga, for two months (December/February). This proved to be beneficial to 
fishermen and the stock and has been retained (Veillon, 1973). The closed season at 
the beginning of the year protects the shrimps when they are particularly small. The 
shore barriers are known locally as 'valakira'. Randrianarijaona and Razafimbelo 
(1983) report that their use is, in fact, illegal but that it has proved difficult to 
prohibit them since many people gain their living in this way. Fishing effort should 
also be limited; two methods are proposed, details of which are given in Marcille 
(1978). 

Le Reste (1978) also recommends that the minimum size limit for the commercial fishery 
should be 10 cm, which is anyway the smallest size of economic value. 

Current legislations are described under the decree of 18 May 1971 (Randrianarijoana and 
Razafimbelo, 1983). Shrimp boats must be licensed; fishing is prohibited from 
15 December to 15 February; the mesh-size of nets is controlled; and boats with motors 
larger than 25 hp may not trawl within two miles of the coast. 

Le Reste (1978) describes the P. indicus fishery in the Baie d'Ambaron in the north-west 
between Cap d'Ambre and Cap St. Andre in detail. The larvae are shed at sea; juveniles 
move to the estuaries and at a size of about 4.5 cm move into the intertidal zone where they 
remain until they reach a size of about 6.5 cm. The annual cycle involves two generations. 
From October to November, a small number of eggs are spawned since the percentage of 
females in the population is low. High temperatures result in favourable conditions for 
juvenile growth. As a result, both the artisanal and commercial fisheries take large catches 
of this generation A between March and June. At this time of year, conditions are less 
favourable for juvenile growth, although there are large numbers of females in the 
population laying eggs, and so the catches of generation B in July and August, taken only 
by the artisanal fishery north of Nosy Be, are small. It is suggested that this fishery could 
be extended further along the coast. It is also recommended that the fishery for generation 
A would ultimately need to be protected from overexploitation although at the time this 
study was carried out (presumably c. 1972), this was thought to be an unlikely occurrence. 



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Fauna 

All coastal stocks have been discovered and in 1973 the majority were already maximally 
exploited, and it was considered that the rest soon would be. About 5000 tonnes were 
produced a year and it was thought that this would increase to 8000 tonnes, but that stocks 
would be unlikely to sustain a higher harvest (Anon., 1973; Marcille and Veillon, 1973a). 
Studies made by the ORSTOM centre at Nosy Be during the 18 years preceding 1973 
showed that there was no hope of developing deep-water trawling on the continental shelf 
as either stocks are poor or the bottom type such as coral makes dredging impossible. 
Further development of the fishery will therefore necessitate the discovery of new stocks, 
possibly on the continental slope. In 1971, ORSTOM therefore started a programme to look 
at the slope between 100-1000 m depth. Details of this study are given in Crosnier and 
Jouannic (1973) and the species are described in Crosnier (1978). 

More than 250 decapod species were caught but probably only a few are of commercial 
value. The following were considered to be of the greatest potential value on account of 
their size and abundance: 

Hymenopenaeus sibogae = Haliporoides sibogae (De Man, 1907). The Jack-knife Shrimp, 
Salicoque Canif; of importance in Japanese fisheries. 

Plesiopenaeus edwardsianus (Johnson, 1868). The Scarlet Shrimp, Gambon Ecarlat; Eastern 
Atlantic where it is fished commercially by Spanish trawlers; the Madagascan form may not 
be the same species. 

Aristaeomorpha foliacea (Risso, 1827). The Giant Red Shrimp, Gambon Rouge. Eastern 
Atlantic, fished by commercial deep sea trawlers off Mediterranean coasts; the taxonomic 
status of the Indo-West Pacific form is not yet clear, and it is not yet fished commercially 
in this area. 

Aristeus virilis (Bate, 1881). The Stout Red Shrimp, Gambon Gaillard. Indo-West Pacific; 
does not yet appear to be fished commercially anywhere 

Aristeus mabahissae Ramadan, 1938. Not yet fished commercially. 

The langoustine Nephrops andamanicus was also considered potentially commercially 
valuable. 

The survey showed that the only suitable area for such a fishery was in the Toliara region 
where stocks were reasonably abundant. Hymenopenaeus sibogae madagascarinum 
and Nephrops andamanicus were found at depths of 425-475 m, and the other species were 
found at depths of 700-800 m. It was recommended that further research should be carried 
out in the Toliara region using a commercial deep-sea fishery vessel rather than a research 
vessel. 

Another small freshwater shrimp species known locally as 'tsivakihiny', Acetes erythraeus 
(Sergestidae), which is widespread from the Red Sea down to South Africa, is fished in the 
muddy, shallow estuaries of the north and north-west coasts by women. The shrimps are 
dried and eaten in coastal villages. The quantities collected are unknown but are reported 
to be large (Crosnier et al., 1962; Le Reste, 1971). 

We are very grateful to Dr Alain Crosnier who provided the information for this section. 



REFERENCES 

Anon. (1973). L'evolution de la peche crevettiere a Madagascar. Bull. Madagascar 

323: 498-506. 
Bardach, J.E., Ryther, J.H. and McLarney, W.O. (1972). Aquaculture. Wiley-Interscience, 

New York and London. 



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Chabanne, J., and Plante, R. (1969). Les populations benthiques (endofaune crevettes 

peneides, poissons) d'une baie de la cdte nord-ouest de Madagascar: ecologie, biologie et 

peche. Cah. ORSTOM, ser. Oceanogr. 7(1): 41-71. 
Chabanne, J. and Plante, R. (1971). Etude des rendements de la peche au chalut des 

crevettes Peneides sur la cote nord-ouest de Madagascar de 1966 a 1970. Doc. sci. Centre 

ORSTOM Nosy-be 23, 19 pp. 
Charbonnier, D. and Crosnier, A. (1961). Quelques donnees sur la peche des langoustes a 

Madagascar. La Peche Maritime 994: 16-18. 
Crosnier, A. (1962). Crustaces Decapodes. Portunidae. Faune de Madagascar 16: 154 pp. 
Crosnier, A. (1965). Crustaces Decapodes. Grapsidae et Ocypodidae. Faune de Madagascar 

18: 143pp. 
Crosnier, A. (1978). Crustaces Decapodes. Peneides Aristeidae (Benthesieyminae, 

Aristeinae, Solenocerinae). Faune de Madagascar 46: 197 pp. 
Crosnier, A. (1983). In litt. 27 April. 

Crosnier, A. and Fourmanoir, P. (1962). La 'Tsivakihiny'. Nat. Malgache 13: 87-88. 
Crosnier, A. and Jouannic, C. (1973). Note d'information sur les prospections de la 

pente continentale malgache effectuees par le N.O. Vauban. Doc. sci. Mission ORSTOM 

Nosy-Be 42: 18 pp. 
Fourmanoir, P. (1935). Le crabe des cocotiers (Birgus latro, famille des Cenobitides). 

Nat. malgache 5(2): 343-344. 
Fourmanoir, P., Crosnier, A. and Charbonnier, D. (1960). Peche a la langouste dans la 

region de Fort-Dauphin (sud-est de Madagascar). Colloque de biologie marine et des 

peches marilimes sur les cotes orientates d'Afrique, Le Cap, Septembre 1960. Doc. mar. 

Biol. 60(4): 5 pp. 
Frontier Albon, D. (1972). Note preliminaire sur un essai de fabrication artisanale de 

NUOC-MAM a partir des residus de l'industrie crevettiere. Doc. sci. Centre ORSTOM 

Nosy-Be 39: 21 pp. 
Holthuis, L.B. (1980). FAO Species catalogue. Vol. 1. Shrimps and Prawns of the World. 

Food and Agriculture Organisation of the United Nations, Rome. 
Le Reste, L. (1970). Contribution a l'etude du rythme d'activite nocturne de Penaeus 

indicus et Parapenaeopsis acclivirostris (Crustacea Decapoda Natantia). Cah. ORSTOM. 

ser. Oceanogr. 8(3): 1-10. 
Le Reste, L. (1971). Rythme saisonnier de la reproduction, migration et croissance des 

postlarves et des jeunes chez la crevette Penaeus indicus H. Milne Edwards en baie 

d'Ambaro, cote nord-ouest de Madagascar. Cah. ORSTOM. ser. Oceanogr. 9(3): 279-292. 
Le Reste, L. (1971). Biologie de Acetes erythraeus (Sergestidae) dans une baie du 

nord-ouest de Madagascar (Baie d'Ambon). Cah. ORSTOM ser. Oceanogr. 8(2): 35-56. 
Le Reste, L. (1973). Etude du recrutement de la crevette Penaeus indicus H. Milne 

Edwards dans la zone de Nosy Be (cote nord-ouest de Madagascar). Cah. ORSTOM, ser. 

Oceanogr. 11(2): 171-178. 
Le Reste, L. (1973). Etude de la repartition spatio-temporelle des larves et jeunes 

postlarves de la crevette Penaeus indicus H. Milne Edwards en baie d'Ambaro (cote 

nord-ouest de Madagascar). Cah. ORSTOM, ser. Oceanogr. 11(2): 179-189. 
Le Reste, L. (1973). Zones de ponte et nurseries de la crevette Penaeus indicus H. Milne 

Edwards le long de la cote nord-ouest de Madagascar. Doc. sci. Centre ORSTOM Nosy Be 

32: 27 pp. 
Le Reste, L. (1978). Biologie d'une population de crevettes Penaeus indicus H. Milne 

Edwards sur la cote nord-ouest de Madagascar. Trav. Doc. ORSTOM 99: 291 pp. 
Le Reste, L., Feno, L. and Rameloson, A. (1976). Etat de nos connaissances sur le crabe 

de vase Scylla serrata Forskal a Madagascar. ORSTOM, Paris. 
Le Reste, L. and Marcille, J. (1973). Reflexions sur les possibility d'amenagement de 

la peche crevettiere a Madagascar. Bull. Madagascar 320: 14-27. 
Le Reste, L. and Marcille, J. (1976). La peche crevettiere a Madagascar. In: Biologie 

marine et exploitation des ressources de V ocean Indien occidental. Communications 

presentees au Colloque Commerson, La Reunion, October 1973, Travaux et Documents de 

I'ORSTOM, No. 47: 221-223. 
Marcille, J. (1972). Les stocks de crevettes peneides cotieres malgaches. Bull. 

Madagascar 311: 387-408. 
Marcille, J. (1978). Dynamique des populations de crevettes peneides exploitees a 

Madagascar. Trav. Doc. ORSTOM 92: 197 pp. 

-104- 



Fauna 

Marcille, J. and Stequert, B. (1974). La peche crevettiere a Madagascar en 1973. 

Evolution des stocks et des pourcentages des differentes especes dans les captures. Doc. 

sci. Mission ORSTOM Nosy-Be, 43: 41 pp. 
Marcille, J. and Veillon, P. (1973a). La peche crevettiere a Madagascar de 1967 a 1972; 

evolution des stocks. Doc. sci. Centre ORSTOM Nosy-Be 35: 28 pp. 
Marcille, J. and Veillon, P.. (1973b). Les stocks de crevettes de Madagascar. Peche 

maritime 1146: 717-720. 
ORSTOM (1973). Publications du Centre ORSTOM de Nosy-be. Liste mise a jour au 

31.12.71. Doc. Sci. Centre ORSTOM Nosy-Be 33: 104 pp. 
ORSTOM (1975). Publications de la Mission ORSTOM de Nosy-Be du 1.1.1972 au 31.12.1974. 

Doc. Sci. Centre ORSTOM Nosy-Be 51: 45 pp. 
Pichon, M. (1964). Contribution a l'etude de 1'ecologie et des methodes de peche des 

Palinuridae dans la region de Nossi-Be. Cah. ORSTOM sir Oceanogr. 2(3): 71-101. 
Ralison, A. (1971). Note sur le stock de crevettes de la cote nord-ouest malgache. 

Service des peches, Majunga, 7 pp. 
Randrianarijaona, P. and Razafimbelo, E. (1983). Rapport national pour Madagascar. 

Report prepared for UNEP Regional Seas East Africa programme. 
Serene, R. (in press). Crustaces Decapodes Brachyoures de l'Ocean Indien Occidental et 

de la Mer Rouge. Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par 

A. Crosnier: Carpiliidae et Menippidae. Faune Tropicale. 
Veillon, P. (1973). Analyse des effets de la fermeture de la peche crevettiere decidee, 

dans certaines zones de Madagascar, du 15 decembre 1972 au 15 fevrier 1973. Doc. sci. 

Centre ORSTOM Nosy-Be 37: 16 pp. 
Warner, G.F. (1977). The Biology of Crabs. Elek Science, London. 



105- 



An environmental profile of Madagascar 

V.ll. OTHER INVERTEBRATES 

Marine 

The marine fauna in general has few endemics since the position of Madagascar in the 
south-west Indian ocean means that most of its fauna is common to the Indo-Pacific with, in 
the south, species from temperate seas (Magnier, 1981). No information is available on the 
conservation status of the following groups. 

a. Corals and other coelenterates 

Two hundred and twenty-two species of octocoralliens have been described from the waters 
around Madagascar, Seychelles, Reunion, Mauritius and Aldabra. Most are common to the 
whole Indo-Pacific; 62 species were known only from the waters of Madagascar and 
neighbouring islands (Tixier-Durivault, 1966). Coral reefs are discussed in detail in Part IV. 

b. Echinoderms 

Thirty species of sea urchin are known but these are rarely eaten by the local people, 
although Heterocentrotus is eaten in the north and Tripneustes gratilla in the south-west. 
Urchins are, however, often used as bait. 

Sea cucumbers are fished for sale and export to the Far East. At the beginning of the 1950s, 
50-70 tonnes were being collected a year (Decary, 1950). Minimum sizes have been established 
for their capture and sale: 1 1 cm for fresh specimens and 8 cm for dried specimens 
(Randrianarijaona and Razafimbelo, 1983). They are collected mainly from the region between 
Androka and the Baie de Mangoky. Recently, it has become apparent that sea cucumbers are 
being over-collected in certain areas, particularly on the south-west coast (Randrianarijaona 
and Razafimbelo, 1983). 

A rich fauna of ophiuroids has been described (Charbonnier and Guille, 1978). 112 species are 
known, including 24 new species and 1 new genus. Most species were found in the littoral. 

c. Ascidians 

Plante and Vasseur (1966) describe three new species but there is no indication that these are 
necessarily endemic. 



Non-marine 

Previously unknown species of hydrozoans, bryozoans and sponges have been found in 
freshwaters but have not yet been fully studied. Rotifers appear to be numerous but have also 
not been studied (Paulian, 1961). 

a. Flatworms 

Large, brightly coloured, black and red flatworms are found in the soil of the rain forests in 
the east (Griveaud 1981). Apart from parasitic forms of Platyhelminthes, two aquatic 
planarians have been described and at least 31 terrestrial planarians, 23 in the genus Bipalium, 
two in Pelmatoplana, six in Rhynchodemidae. An unpigmented planarian was found in the 
important cave of Andranoboka, but in general planarian diversity seems to be low (Paulian, 
1961). 

b. Nemertines 

At least one species is known (Paulian, 1961). 



■106- 



Fauna 



c. Annelida 



Earthworms (Hakatany or Kankana) can reach large sizes, up to 75 cm long (Decary, 1950). 
Leeches are known as 'linta', 'dinta' or 'ditamaka' (Decary, 1950). Small forest leeches are 
abundant in the soil and foliage in the rain forests in the east (Griveaud, 1981). A species 
of Philaemon and three species in the genus Haemadipsa are known, from the rain forests of 
the east (Paulian, 1961). 



REFERENCES 

Charbonnier, G. and Guille, A. (1978). Echinodermes: Ophiurides. Faune de Madagascar 

48: 248 pp. 
Decary, R. (1950). La faune malgache, son role dans les croyances et les usages 

indigenes. Payot, Paris. 236 pp. 
Griveaud, P. (1981). Les invertebres: un univers prodigieux. Ch. 3. In: Oberle, P. 

(Ed.), Madagascar: un sanctuaire de la nature. Lechevalier, S.A.R.L., Paris. 
Magnier, Y. (1981). Merveilles sous-marines. Ch. 7. In: Oberle, P. (Ed.), Madagascar: 

un sanctuaire de la nature. Lechevalier, Paris. 
Paulian, R. (1961). La zoogeographie de Madagascar et des iles voisines. Faune de 

Madagascar 13: 1-485. 
Plante, R. and Vasseur, P. (1966). Sur une collection d'Ascidies de la region de Tulear 

(cote sud-ouest de Madagascar). Ann. Univ. Mad. 4: 143-159. 
Randrianarijaona, P. and Razafimbelo, E. (1983). Rapport national pour Madagascar. 

Report prepared for UNEP Regional Seas East Africa programme. 
Tixier-Durivault, A. (1966). Octocoralliaires. Faune de Madagascar 21: 456 pp. 



■107- 



PART VI. PROTECTED AREAS 
VI. 1. INTRODUCTION 

Legislation 

Five different categories of protected area are recognised: 

a. Reserves Naturelles Integrates (Strict Nature Reserves). 

b. Pares Nationaux (National Parks). 

c. Reserves Speciales (Special Reserves) 

d. Forets Classees (Classified Forests). 

e. Perimetres de Reboisement et de Restauration (Reafforestation Zones). 

a. Legislation for the Strict Nature Reserves is chiefly contained in Decree 66-242 (01.06.66); 
the network of reserves was originally set up in 1927. These reserves are part of the national 
forest domain; access is strictly forbidden, other than for scientific research purposes (which in 
each case must be authorised by the Ministry in charge of the Direction des Eaux et Forets). 
Each reserve comprises a station for a deputy forester and each station is divided into two or 
three sectors under the responsibility of auxiliaries. At present there are 1 1 Strict Natu r e 
Reserves; a twelfth, R.N.I. No 2 on the Masoala Peninsula, was de-gazetted by Decret 64-381 
of 16.09.64. and is now a Classified Forest. 

b. National Parks legislation is contained in Decrees 58-07 (28.10.58) and 62-371 (19.07.62). 
Access to the Parks is controlled. Rights are accorded to neighbouring villagers for the 
exploitation of certain forest products; these rights are tied to duties incumbent on the 
beneficiaries. There are two national parks at present. 

c. Special Reserves have been set up by a series of decrees and are designed to protect certain 
animal or plant species. Access to such reserves is free, but hunting, fishing, pasturing of 
livestock, collection of natural products and introduction of vegetation or animal species are all 
forbidden; however, these areas do not generally have supervisory personnel. There are 23 
Special Reserves, of which the most recently created (November 1985) is Beza Mahafaly in 
Toliara Province. 

d. Classified Forests are the subject of individual ministerial decrees; points in common are: 

- all forest exploitation is forbidden; 

- they are intended to constitute 'forest reserves' in the economic sense of the term; 

- local inhabitants can exercise certain traditional rights of usage (limited to the 
collection of minor forest products such as honey and raffia); 

- classified forests are created under an initiative from the Minister in charge of the 
Direction des Eaux et Forets on the advice of a commission composed of representatives 
of the administration and local inhabitants; protection is not necessarily permanent. 

e. Reafforestation and Restoration Zones are set up following the same procedure as for 
Classified Forests above. Their aim is principally stabilisation and protection of watersheds, 
and prevention of erosion. Land-use is regulated within these regions, and they benefit from 
management practices such as reafforestation, management of pasture and use of antierosion 
measures and devices. 

The original definitions for parks and reserves were based on the 1933 London Convention 
(which was accepted in Malagasy law on 25 January 1937), and the African Convention for the 
Conservation of Nature and Natural Resources (1968) is invoked for the protection and 
conservation of all protected areas. 

Administration 

This is the responsibility of the Administration Forestiere within the Direction des Eaux et 
Forets which at the present time comes under the Ministere de la Production Animale (Elevage 



109- 



An environmental profile of Madagascar 

et Peche) et des Eaux et Forets. The Direction des Eaux et Forets is also responsible for forests 
and freshwater fisheries, while other departments within the same Ministry deal with marine 
fisheries and agriculture. 



Total area under protection 



National Parks 

Isalo (No.2) 

Montagne d'Ambre (No.l) 



Total 



Nature Reserves 

Tsingy de Bemaraha (No.9) 
Andohahela (No.l 1) 
Zahamena (No. 3) 
Ankarafantsika (No. 7) 
Marojejy (No. 12) 
Tsaratanana (No.4) 
Tsimanampetsotsa (No. 10) 
Andringitra (No. 5) 
Tsingy de Namoroka (No. 8) 
Betampona (No.l) 
Lokobe (on Nosy Be) (No. 6) 

Total 

Special Reserves 

Ambatovaky (Toamasina Province) 

Marotandrano (Mahajanga) 

Manongarivo (Antseranana) 

Analamerana (Antseranana) 

Anjanaharibe-Sud (Antseranana) 

Kalambatritra (Fianarantsoa) 

Ambohijanahary (Mahajanga) 

Kasijy (Mahajanga) 

Ankara (Antseranana) 

Tampoketsa d'Analamaitso (Mahajanga) 

Bemarivo (Mahajanga) 

Maningozo (Antseranana) 

Andranomena (Toliara) 

Ambohitantely (Antananarivo) 

Manombo (Fianarantsoa) 

Foret d'Ambre (Antseranana) 

Bora (Mahajanga) 

Pic d'lvohibe (Fianarantsoa) 

Cap Sainte Marie (Toliara) 

Perinet-Analamazoatra (Toamasina) 

Mangerivola (Toamasina) 

Beza Mahafaly (Toliara) 

Nosy Mangabe (Toamasina) 

Total 

Classified forests 

There are 158 of these with total area 

Reafforestation areas 

There are 77 of these covering 



(hectares) 



81 540 


18 200 


99 740 


152 000 


76 020 


73 160 


60 520 


60 150 


48 622 


43 200 


31 160 


21 742 


2 228 


740 


569 542 


60 050 


42 200 


35 250 


34 700 


32 100 


28 250 


24 750 


18 800 


18 220 


17 150 


11 570 


7 900 


6 420 


5 600 


5 020 


4 810 


4 780 


3 450 


1 750 


810 


800 


600 


520 


365 500 


ca 2 671 000 


ca 823 978 



110- 



Protected areas 

Finally, a decree of 23 May 1923 declared the following as protected areas for the 
turtles Chelonia mydas and 'Chelonia imbricata' (=Eretmochelys imbricata): Nosy 
Ovambo; Nosy Iranja, Chesterfield Island, Nosy Trozona, Nosy Ve and Europa (this last 
is now under French control). 

Information sheets 

The following section provides information on the two national parks, the eleven existing 
natural reserves, four special reserves (Ambohitantely, Beza Mahafaly, Nosy Mangabe and 
Perinet-Analamazoatra) and the private reserves at Berenty and Analabe. Format and contents 
are modified versions of those provided in the forthcoming IUCN Directory of Afrotropical 
Protected Areas. 

A preliminary faunal list for each protected area is provided. It should be stressed that these 
lists are in large part incomplete, and are more a reflection of the available information for 
different faunal groups in each of the areas than of their relative faunal diversity. In general 
the birds and primates are the best-known components of the fauna, although for some areas 
(e.g. R.N.I, de Tsingy de Bemaraha (No.9), the largest reserve in Madagascar) information is 
still very incomplete even for these. For other groups, good information is available for only a 
very few protected areas. For some sites, records have been included if these refer to the 
general area of the reserve (e.g. Massif d'Andringitra (R.N.I. No. 5), Montagne d'Ambre (Pare 
National No.l), Massif de Tsaratanana (R.N.I. No.4)), though it is possible that in some of 
these cases the records are from outside the reserves themselves. For a few cases, notably 
Lokobe (R.N.I. No. 6) and Andohahela (R.N.I. No.ll), some records have been included if they 
refer to the more extensive areas in which the reserves are situated (Nosy Be island and the 
Anosyennes hills respectively), though this is made clear in each case. 

In general, only endemic species have been included; where exceptions have been made these 
are clearly indicated (birds endemic to the Malagasy region but not confined to Madagascar are 
denoted (R)). 

Species which are believed on current knowledge to be restricted to a given reserve or reserve 
area (see above) are marked *; doubtful records are indicated with '?'. 

Much of the information quoted has been provided by recent observers; in particular we are 
extremely grateful to O. Langrand, M. Nicoll, S. O'Connor and M. Pidgeon for access to their 
extensive unpublished data, mainly on birds and mammals, and to Q Bloxam, D. Curl and 
C. Raxworthy for providing information on the herpetofauna of several of the protected areas. 
Most of the remaining records are derived from previously published data; further information 
and references for these are provided in Part V and Appendices 2 and 3. It should be noted 
that many of the published records, particularly those for the rarer bird species, are relatively 
old and may not have been re-confirmed more recently. 

PRINCIPAL REFERENCES 

Anon. (Direction des Eaux et Forets et de la Conservation de Sols, Ministere de 

l'Agriculture, de l'Expansion Rurale et du Ravitaillement) (1969). Domaine Forestier 

1969. Monuments naturels et sites. Madagascar et Comores. 52 pp. 
Andriamampianina, J. (1981). Les reserves naturelles et la protection de la nature a 

Madagascar. In: Oberle, P. (Ed.) Madagascar, un sanctuaire de la nature. Lechevalier 

S.A.R.L., Paris. 
Andriamampianina, J. (1984). Traditional land use and nature conservation in Madagascar. 

Workshop paper. World congress on National Parks, Bali, Indonesia, 11-22 October 1982. 
Conference de Tananarive (1972). Comptes rendus de la Conference Internationale sur la 

Conservation de la Nature et de ses Ressources a Madagascar. Tananarive, Madagascar. 

7-11 October 1970. Publication IUCN Nouvelle Series Document supplementaires No. 36. 
Griveaud, P. and Albignac, R. (1972). The problems of nature conservation in Madagascar. 

In: Battistini, R. and Richard-Vindard G. (Eds) Biogeography and Ecology of 

Madagascar. Monographiae Biologicae 21. Junk, the Hague. 
IUCN/WWF Project 1968. Promotion of conservation in Madagascar. 
IUCN/WWF Project 1951. Provision and equipping of a mobile education unit. 

-Ill- 



An environmental profile of Madagascar 

IUCN/WWF Project 1952. Establishment, promotion and equipment for reserves and parks. 
Jolly, A., Oberle, P. and Albignac, E.R. (Eds) (1984). Key Environments - Madagascar. 

Pergamon Press, Oxford. 
Martin, C. (1982). Rapport de la mission technique WWF/IUCN a Madagascar 1981. 

IUCN/WWF, Gland. 
Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF - Protected areas 

programme in Madagascar. Unpd. report, 62 pp. 
Randrianarijaona, P. and Razafimbelo, E. (1983). Inventaire des Ecosystemes: Rapport 

National pour Madagascar. Report to IUCN, UNEP Regional Seas Programme for East 

Africa. 



NAME Pare National de la MONTAGNE D'AMBRE (No.l) 

MANAGEMENT CATEGORY II (National Park). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 28 October 1958 by Decree No. 58-07. 

GEOGRAPHICAL LOCATION 12°28'-44'S, 49°04'-13'E. At the northern point of 

Madagascar near the town of Antseranana. 

ALTITUDE 850-1474 m. 

AREA 18 200 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The park lies within a volcanic massif which is composed essentially 
of basaltic rock of origin ca 14 myBP., and consists of a line of summits some 30 km long, 
oriented north-south, the tallest being the Pic d'Ambre (1475 m). The park contains a crater 
lake and cascades, and is a place of great beauty. The region of Antseranana has a tropical 
climate with a marked dry season from May to December during which a strong south-east 
trade wind blows (the 'Varatraza') and a wet season from January to April with less than 
1000 mm rainfall. The Montagne d'Ambre, however, has a microclimate characteristic of the 
eastern rainforest with very high rainfall (3585 mm per year has been measured) throughout 
the year, with a maximum in January - February. The area thus serves as an important rain 
catchment area for Antseranana. 

VEGETATION Vegetation is principally upland tropical moist forest, with trees reaching 
30 m height and including species such as Canarium madagascariense, Gluta tourtour, 
Terminalia mantali and Dalbergia sp.; palms (e.g. Neodypsis and Chrysalidocarpus spp.) and 
tree-ferns (Cyathea spp.) are abundant in the understorey and there is a very diverse epiphytic 
flora, including ferns (e.g. Platycerium. Drynaria and Asplenium) orchids (e.g. Bulbophyllum, 
Angraecum and Aeranthes) and lichens. Around the Station Forestiere des Roussettes there are 
plantations of introduced species including pines, Eucalyptus and Araucaria. 

ZONING None. 

CONSERVATION MANAGEMENT The management of the park is the responsibility of the 
Circonscription Forestiere at Antseranana. In 1985, 23 km of the park boundary were 
re-cleared - previously the park's limits had been badly marked on the ground; the path giving 
access to the park was renovated, as was the 'gite d'etape' which provides accommodation at 
the Station Forestiere des Roussettes situated on the northern boundary of the park (Langrand 
and Lenormand, 1985). Reafforestation work has been undertaken at the park's boundaries by 
an association for nature conservation set up be teachers in Antseranana in 1983 (ASSE), in 
collaboration with Eaux et Forets agents based at the Station Forestiere des Roussettes. 

-112- 



Protected areas 

DISTURBANCES OR DEFICIENCIES In general the park is inadequately guarded - there is 
only one agent who lives too far from the park (in Antseranana) to be able to carry out 
effective surveillance. Bush fires threaten the edges of the national park every year, and 
destroy eucalyptus plantations growing at the boundaries; such fires are usually the result of 
uncontrolled pasture burning. Illegal forest exploitation takes place in areas where access by 
Eaux et Forets agents is difficult, particularly in the south. Collection of ornamental plants 
such as orchids, palms, and arborescent ferns for sale in Antseranana is a problem as is 
poaching, particularly of lemurs (Lemur coronatus, L. fulvus) and the Crested Ibis Lophotibis 
cristata; such poaching was said in 1986 to be widespread and increasing. Livestock range 
freely, though in relatively small numbers, through the park. 

SCIENTIFIC RESEARCH There have been numerous collecting trips to the area since 1893. 

VISITOR AND SCIENTIFIC FACILITIES Access is relatively easy from Antseranana (50 km 
north of the park) along an asphalt road. There are some 30 km of paths within the park, but 
their state of repair varies greatly from year to year. The 'jardin botanique' path passes 
numerous different trees that have been identified and labelled for visitors. There is 
accomodation in the form of a 'gite d'etape' at the Station Forestiere des Roussettes (Langrand 
and Lenormand, 1985). 

PRINCIPAL REFERENCE MATERIAL 

Langrand, O. and Lenormand, B. (1985). Presentation sommaire du Pare National de la 

Montagne d'Ambre. Unpd. report, 9 pp. 
Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF - Protected areas 

programme in Madagascar. Unpd. report, 62 pp. 

STAFF One Eaux et Forets agent. 

BUDGET Salaries paid by the government. WWF Direction a Madagascar has provided 
material and financial support for the park. 

LOCAL PARK OR RESERVE ADMINISTRATION Headquarters is at Roussettes. 

FAUNA 



Birds 

Tachybaptus pelzelnii 

Ardeola idae (R) 

Lophotibis cristata 

Anas bernieri 

Haliaeetus vociferoides 

Aviceda madagascariensis 

Accipiter francesii (R) 

Buteo brachypterus 

Falco newtoni (R) 

Margaroperdix madagarensis 

Turnix nigricollis 

Dryolimnas cuvieri 

Sarothrura insularis 

Streptopelia picturata (R) 

Treron austral is (R) 

Alectroenas madagascariensis 

Coracopsis vasa (R) 

Coracopsis nigra (R) 

Coua cristata 

Cuculus rochii (R) 

Centropus toulou (R) 

Otus rutilus (R) 

Asio madagascariensis 

Caprimulgus madagascariensis (R) 



Zoonavena grandidieri (R) 
Alcedo vintsioides (R) 
Ipsidina madagascariensis 
Eurystomus glaucurus (R) 
Atelornis pittoides 
Leptosomus discolor (R) 
Mirafra hova 
Motacilla flaviventris 
Coracina cinerea (R) 
Phyllastrephus madagascariensis 
Phyllastrephus zosterops 
Hypsipetes madagascariensis (R) 
Calicalicus madagascariensis 
Vanga curvirostris 
Leptopterus viridis 
Leptopterus chabert 
Leptopterus madagascarinus (R) 
Copsychus albospecularis 
Monticola sharpei 
Neomixis tenella 
Nesillas typica 
Cisticola cherina 
Newtonia amphichroa 
Newtonia brunneicauda 



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An environmental profile of Madagascar 



Terpsiphone mutata (R) 
Oxylabes madagascariensis 
Nectarinia souimanga (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Lonchura nana (R) 

Mammals 

Lemur coronatus 
Lemur fulvus 
Lepilemur septentrionalis 
Phaner furcifer 
Cheirogaleus major 
Microcebus rufus 



Ploceus nelicourvi 

Foudia madagascariensis (R) 

Foudia omissa 

Saroglossa aurata 

Dicrurus forficatus (R) 



Cryptoprocta ferox 
Galidia elegans 
Tenrec ecaudatus 
Setifer setosus 
Pteropus rufus 



In addition Microgale prolixacaudata, M. parvula and M. drouhardi have locality records of 
'Antseranana' and probably occur in the national park. 



Amphibians 

Cophyla phyllodactyla 
Mantidactylus femoralis 
Mantidactylus granulatus 
Mantidactylus ulcer osus 



*Mantipus laevipes 
Mantidactylus flavicrus 
Mantidactylus lugubris 



Reptiles 

Homopholis boivini? 
*Uroplatus alluaudi 
*Brookesia tuberculata 

Amphiglossus ardouini 
*Paracontias brocchi 
*Pseudoxyrhopus ambreensis 

Acrantophis madagascariensis 

Sanzinia madagascariensis 



Lygodactylus madagascariense 
Uroplatus ebenaui 
Chamaeleo pardalis 
*Androngo allaudi 
Alluaudina belly i 
Leioheterodon madagascariensis 
Ithycyphus miniatus 
Liophidium rhodogaster 



Nonmarine Molluscs 

Tropidophora alluaudi 
Tropidophora deliciosa 

*Tropidophora propeconsocia 
Tropidophora surda 
Tropidophora vignali 
Tropidophora winckworthi 

*Kalidos anceyanus 
Kalidos andapaensis 

*Kalidos dautzenbergianus 

*Kalidos fallax 

* Kalidos glessi 
Kalidos humbloti 



*Kalidos tenebricus 
*Ampelita alluaudi 

Ampelita atropos (foret des Rousettes) 

Ampelita dingeoni 
* Ampelita lamothei 
* Ampelita subatropos (foret des Rousettes) 

Clavator moreleti 

Edentulina alluaudi (foret des Rousettes) 

Edentulina nitens 

Macrochlamys stumpfii 

Edouardia rufoniger 



Nonmarine Crustacea 
Isopoda: 

Suarezia heterodoxa 
Decapoda: 

Hydrothelphusa agilis madagascariensis 

Lepidoptera 
Papilionidae 

Graph ium endochus 

Danaidae 

Amauris nossima dijuncta 



Nymphalidae 

Charaxes andranodorus 
Hypolimnas dexithea 
Cymothoe lembertoni 



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Protected areas 

NAME Pare National de 1'ISALO (No.2) 

MANAGEMENT CATEGORY II (National Park). 

LEGAL PROTECTION Total, except that inhabitants of Ranohira are authorized to harvest 
silkworm cocoons (Bocerus sp.) and Tapia Uapaca bojeri. 

DATE ESTABLISHED 19 July 1962 by Decree No. 62-371. 

GEOGRAPHICAL LOCATION West of Ihosy, in the province of Fianarantsoa. 22°10'-41'S, 
49°10'-21'E. 

ALTITUDE 514-1268 m. 

AREA 81 540 ha. 

LAND TENURE Central Government. 

PHYSICAL FEATURES The park encompasses the whole of the sandstone Isalo Massif and 
has very varied relief. The south and east of the massif has characteristically ruiniform relief 
ranging in altitude from 820 to 1240 m above sea level, comprising 100-200 m deep, 
narrow-floored canyons extending for several kilometres. These have temporary or permanent 
watercourses. The northern and western parts of the massif are not ruiniform but also have 
200-300 m high cliffs enclosing narrow, deep gorges. There are clear, fast-flowing streams 
which join the Malio River to the west and the Menamaty in the east, both these being 
tributaries of the Mangoky. Climate is dry and tropical, with a rainfall of ca 850 mm. A 
marked dry season occurs between June and August; the wet season runs from October to 
March with highest rainfall recorded in November and December. Average monthly 
temperatures are lowest in June (17.1°C) and highest in February (25.rC). Winds (velocity 
6-12 m/s) blow throughout the year. 

VEGETATION Much of the vegetation is very degraded. A dry deciduous forest covers ca 
20% of park, largely in the north and west (Sahanafo, Ankikiky and Angodongodona). In some 
areas the fire-resistant Uapaca bojeri is the only canopy species. Canyon floors with 
watercourses support evergreen forest with Eugenia, Pandanus, Chrysalidocarpus and two fern 
species predominating. Rock-dwelling vegetation is very abundant and principally consists of 
mPachypodium rosulatum and Aloe isaloensis, both endemic to the massif. 

ZONING None. 

CONSERVATION MANAGEMENT None at present. 

DISTURBANCES OR DEFICIENCIES Bush fires consitute the most serious threat to the 
park: the pastures bordering the park are subject to annual uncontrolled burning which spreads 
into the surviving wooded areas. Only isolated areas such as canyons are protected from fire. 
The ranging of Zebus within the park has been a problem, though in 1986 was noted to have 
considerably decreased over the past few years owing to the extent of cattle-rustling in the 
region. People living around the park often enter to collect fuel wood {Uapaca) and building 
wood (e.g. Weinmannia), and to hunt. Poaching principally involves lemurs {Lemur catta), the 
bush pig Potamochoerus larvatus, guinea fowl Numida meleagris, fruit bats {Pteropus rufus 
or Eidolon helvum) and eels. Honey is also collected within the park. 

SCIENTIFIC RESEARCH A historical study of the rock shelter known as the 'Grotte des 
portugais' at Tenika was carried out in 1960-62; despite its name, it is believed likely to be of 
11th century Arab origin. 

VISITOR AND SCIENTIFIC FACILITIES None, although the park is run as a tourist 
attraction by the agents of the Direction des Eaux et Forets. Visitors (82 in 1985, 67 in 1986) 
pay an entrance fee of 1500 F.M.G. (which goes direct to the local community, not to the 
Direction des Eaux et Forets) and are guided by the Eaux et Forets agents or other local 

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guides. Ranohira, the main town bordering the reserve, lies on the Route Nationale linking 
Toliara to Fianarantsoa; it has shops and a hotel. 

PRINCIPAL REFERENCE MATERIAL 

Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF - Protected areas 

programme in Madagascar. Unpd. report, 62 pp. 
Revue de l'Office du Tourisme de Madagascar No. 33 (La Grotte des Portugais by P. Ginther 

and V.C. Herbert). 

STAFF One agent and one auxiliary. 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Administration is at Ranohira. 

FAUNA 

Birds 

The following species have been recorded by O. Langrand (unpublished data, in litt., 
22.12.86). 



Tachybaptus pelzelnii 
Buteo brachypterus 
Falco newtoni (R) 
Margaroperdix madagarensis 
Turnix nigricollis 
Pterocles personatus 
Streptopelia picturata (R) 
Coracopsis nigra (R) 
Agapornis cana 
Cuculus rochii (R) 
Centropus toulou (R) 
Ninox superciliaris 
Caprimulgus madagascariensis (R) 
Zoonavena grandidieri (R) 
Alcedo vintsioides (R) 
Eurystomus glaucurus (R) 

Mammalia 

Propithecus verreauxi 
Lemur catta 
Lemur fulvus 

Reptiles 

Acrantophis dumerilii 



Leptosomus discolor (R) 
Mirafra hova 
Phedina borbonica (R) 
Motacilla flaviventris 
Coracina cinerea (R) 
Leptopterus viridis 
Copsychus albospecularis 
Monticola bensoni 
Nesillas typica 
Cisticola cherina 
Newtonia brunneicauda 
Neomixis tenella 
Terpsi phone mutata (R) 
Nectarinia sow manga (R) 
Foudia madagascariensis (R) 
Dicrurus forficatus (R) 



Cryptoprocta ferox 
Tenrec ecaudatus 
Tenrec setifer 






Oplurus sp. 



NAME Reserve Naturelle Integrate de BETAMPONA (No.l). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927. The area of the reserve was fixed by Decree 
66-242 of June 1966. 

GEOGRAPHICAL LOCATION 17°51'-55'S, 49 12'-15'E. The reserve is situated 40 km to 
the north-west of Toamasina in Toamasina Province. 



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Protected areas 

ALTITUDE 275-650 m. 

AREA 2228 ha. 

LAND TENURE State land. 

PHYSICAL FEATURES The reserve occupies a rocky spur which dominates the coastal plain 
and consists of a family of ridges varying in height from 320 m to 650 m above sea level. It is 
a rugged zone where numerous streams have their sources. Climate is humid tropical with 
precipitation over 2000 mm a year and no dry months. Mean annual temperature is between 
21° and 24°C, though in the colder months mean temperature can be as low as 12°C. The 
reserve is an example of the natural low altitude biotope of the eastern region of Madagascar. 

VEGETATION Betampona is the sole forested massif in a vast deforested area, and although 
the sides of the massif have been deforested, the vegetation has re-established itself to some 
degree (see below). The undegraded vegetation is low altitude dense evergreen rain forest 
which is extremely rich floristically and is the type locality for many species. It is 
characterised by species of Myristicaceae and Anthostema (Euphorbiaceae); other notable 
species are Canarium madagascariensis (Burseraceae), Sideroxylon sp., Faucherea ursii (both 
Sapotaceae), Rhopalocarpus 

sp. (Sphaerosepalaceae), Hirtella sp. (Chrysobalanaceae). Also present are members of the tribe 
Areceae (Palmae) and the families Rubiaceae, Araliaceae, Ebenaceae, Sapindaceae, Loraceae, 
Myrtaceae, Flacourtiaceae and Leguminosae. Local dominance of Uapaca thouarsii and glades 
of the bamboo Cephalostachyum madagascariensis occur. Much of the central southern edge 
of the reserve, notably along the Fontsiamavo River and its tributaries, is composed of the 
colonising Eugenia jambos. Secondary forest has considerably advanced, especially along the 
central track from Sorintsandry to Marovato, an area which had not been cleared in 1947. 

ZONING There was formerly a 200 m protective zone around the reserve, set up in 1935; 
Pollock reported in 1985 that it had recently been reclaimed by local villagers for residential 
and exploitative purposes. 

CONSERVATION MANAGEMENT The reserve is clearly delimited by a 3-metre wide band 
cleared around the perimeter and has a good internal path network. Insufficient funds and 
staff are available at present for further management. 

DISTURBANCES OR DEFICIENCIES A degraded zone some 1000-1500 m wide lies within 
the reserve boundaries, consisting largely of 20-25 year old growth of Ravenala 
madagascariensis and other 'savoka' species. This represents an area of regeneration following 
cultivation: the existence of tavy cultivation and the corresponding threat to the forest edges 
was first remarked in 1908. The situation had not markedly improved in 1931 and although 
the Reserve Integrate was first declared in 1932 and the protective zone in 1935, real protection 
was not initiated until 1949; the regeneration presumably dates from then. Forest cover within 
the reserve thus totals some 1000 ha. It is surrounded by numerous villages, situated within 
1 km of the reserve and often less, and there are incursions into the reserve: Pollock noted in 
1984 that there were clear signs of crayfish catching in the upper Fontsiamavo River within the 
reserve, and that a local merchant had recently been arrested for shooting lemurs in the central 
part of the reserve. 

SCIENTIFIC RESEARCH None appears to have been carried out recently. 

VISITOR AND SCIENTIFIC FACILITIES Access to the reserve is on foot (90 minutes walk) 
from the village of Fontsimavo, which is 90 minutes by car from Toamasina (Pollock, 1985). 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

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An environmental profile of Madagascar 

Pollock, J.I. (1985). Preliminary report on a mission to Madagascar by Dr. J.I. Pollock 
in August and September 1984. Unpublished report, 10 pp. 

STAFF One agent and two auxiliaries, full-time. 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Headquarters at Rendrirendry. This 
village consists of the dwelling quarters of two families, the local office of the Direction des 
Eaux et Forets and a 'gite d'etape' or rest-post for the guardian of the reserve, who lives in 
Toamasina. 

FAUNA 



Birds 

Lophotibis cristata 
Treron australis 
Alectroenas madagascariensis 
Ipsidina madagascariensis 
Leptosomus discolor 

Mammals 

Avahi laniger 
Hapalemur griseus 
Cheirogaleus major 
Lemur fulvus 
Lepilemur mustelinus 
Phaner furcifer 
Cryprotprocta ferox 
Eupleres goudotii 



Philepitta castanea 
Vanga curvirostris 
Terpsiphone mutata 
Zosterops maderaspatana 



Varecia variegata 

Microcebus rufus 

Indri indri 

Propithecus diadema (possibly) 

Daubentonia madagascariensis 

Galidia elegans 

Fossa fossa 

Galidictis striata (reported) 



Amphibians 

* Plelhodontohyla coudreaui 

Reptiles 

Ebenavia inunguis 

Nonmarine Molluscs 

Ampelita lanx 
A. sepulchralis 
Tropidophora tricarinata 

Nonmarine Crustacea 
Decapoda: 

Hydrothelphusa agilis madagascariensis 



Helicophanta magnifica 
Macrochlamys stumpfii 
Kalidos bournei 



NAME Reserve Naturelle Integrate de ZAHAMENA (No. 3). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927. 

GEOGRAPHICAL LOCATION 17°26'-44'S, 48°56'-49 o 00'E; east of Ambatondrazaka in 
Toamasina Province, bordered to the south by the Onibe, to the east by the Ihofika River, to 
the west by the Vohimahery and to the north by the track from Sahatavy to Imerimandroso. 

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Protected areas 

ALTITUDE 500-1500 m. 

AREA 73 160 ha, essentially divided into two blocks. 

LAND TENURE State land. 

PHYSICAL FEATURES The reserve consists of two zones well separated by a large piece of 
enclosed land. Relief is very uneven with steep-sided valleys; large landslides were recorded in 
the 1950s. Climate varies with altitude from an equatorial or tropical hurnid type, to one with 
a two or three month dry season (August to October). Mean annual rainfall is 1500-2000 mm 
but may be considerably higher in the east. Average temperature of the coolest month is 
between 10° and 15°C at higher altitudes. 

VEGETATION Generally characteristic of the eastern escarpment, with primary and 

secondary tropical evergreen forest, changing from forest with Myristicaceae and Anthostema 
typical at low altitude to forest with Tambourissa and Weissmannia between 800 and 1300 m. 
With increasing altitude, plants with temperate affinities appear, and there are notable 
deciduous species and belts of bamboo; the higher altitude forest when degraded transforms 
into ericaceous scrub with Philippia, Agauria and Helichrysum. 

ZONING None. 

CONSERVATION MANAGEMENT Relatively little at present as the reserve is undermanned; 
the 1985 University of London Expedition financed the construction of a 2 km firebreak along 
the western edge of the reserve. 

DISTURBANCES OR DEFICIENCIES Most of the western half and south-eastern parts of 
the reserve were noted in 1985 as having an unbroken canopy of good forest; a trail through 
the the south-centre of the reserve passed through an abandoned clearing which was reverting 
to forest. Other than this, only a few trees had been selectively felled. However, the 
north-eastern corner of the reserve (near the enclave - see below) was under greater threat: the 
trail there was heavily used, an area of ca 1 sq. km had been cleared for tavy and three lemur 
traps were found. The western edge of the reserve adjoined eucalyptus plantations and 
cultivated land, and faced threats from logging and fire. The central enclave was part of the 
reserve as it was originally gazetted; the continued presence of several villages there led to the 
area being degazetted in 1966 (Decree 66-242). These villages were reported in 1971 as 
growing in size and representing a long-term threat to the integrity of the reserve; cultivation 
(by tavy), livestock grazing and poaching apparently occurred around the edges of the enclave 
(in the north-east corner of the reserve). In 1985, it was noted that the enclave had been 
cleared for agriculture except for some secondary scrub and a tiny area of relict forest. 

SCIENTIFIC RESEARCH Important entomological work has been conducted in the area. In 
1985 a five person expedition from the University of London spent 32 days in the reserve, 
primarily surveying the avifauna, though also collecting amphibians and making observations 
on mammals and plants (particularly pteridophytes) (Thompson et al., in press). 

VISITOR AND SCIENTIFIC FACILITIES None. Much of the reserve is virtually 

impenetrable and lacks trails. 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

Thompson, P.M., Raxworthy, C.J., Murdoch, D.A., Quansah, N. and Stephenson, P.J. (in 
press). Zahamena Forest (Madagascar) Expedition, 1985. ICBP Study Report. ICBP, 
Cambridge. 

STAFF One agent and two auxiliaries full-time. 



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An environmental profile of Madagascar 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Headquarters at Manakambahiny Est. 



FAUNA 

Birds 

The 1985 University of London expedition to Zahamena recorded the following endemic 
birds. Those in parentheses were only recorded in the enclave, not in the reserve proper, but 
many, if not all, are thought likely to occur in the reserve. 



Polyboroides radiatus 

Accipiter madagascariensis 

(Accipiter francesii (R)) 

Buteo brachypterus 

Falco newtoni (R) 

Dryolimnas cuvieri 

Streptopelia picturata (R) 

Alectroenas madagascariensis 

Coracopsis nigra (R) 

Coracopsis vasa (R) 

Coua serriana 

Coua reynaudii 

Coua caerulea 

Centropus toulou (R) 

Otus rutilus (R) 

(Caprimulgus madagascariensis (R)) 

Zoonavena grandidieri (R) 

Alcedo vintsioides (R) 

Ipsidina madagascariensis 

Brachypteracias leptosomus 

Leptosomus discolor (R) 

Philepitta castanea 

Neodrepanis coruscans 

Phedina borbonica (R) 

Coracina cinerea (R) 

Phyllastrephus madagascariensis 

Phyllastrephus zosterops 

Hypsi petes madagascariensis (R) 

Tylas eduardi 

Lophotibis cristata has been reported to occur 
1985 expedition. 

Mammals 

Microcebus rufus 
Cheirogaleus major 
Avahi laniger 
Indri indri 
Propilhecus diadema 
Daubentonia madagascariensis 



Calicalicus madagascariensis 
Leptopterus chabert 
Leptopterus viridis 
Leptopterus madagascarinus (R) 
Oriolia bernieri 
Euryceros prevostii 
Hypositta corallirostris 
Copsychus albospecularis 
Neomixis tenella 
Neomixis viridis 
Neomixis striatigula 
Hartertula flavoviridis 
Oxylabes madagascariensis 
Nesillas typica 
Cisticola cherina 
Randia pseudozosterops 
Newtonia amphichroa 
Newtonia brunneicauda 
Pseudobias wardi 
Terpsiphone mutata (R) 
Nectarinia souimanga (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Lonchura nana (R) 
Ploceus nelicourvi 
(Foudia madagascariensis (R)) 
Foudia omissa 
(Saroglossa aurata) 
Dicrurus forficatus (R) 

in the reserve but was not observed by the 



Hapalemur griseus 
Lemur fulvus 
Lemur rubriventer 
Lepilemur mustelinus 
Varecia variegata 
Galidia elegans 



Amphibians 

The following species were collected by the 1985 University of London Expedition. 



Ptychadena mascariensis 
Aglyptodactylus madagascariensis 
Mantidactylus guttulatus 
Mantidactylus majori 
Mantidactylus wittei 



Mantidactylus pliciferus? 
Mantidactylus aerumnalis 
Mantidactylus betsileanus 
Boophis viridis? 
Mantella cowani 



120- 



Protected areas 



Mantidactylus femoralis 
Mantidactylus luteus 

Reptiles 

Phelsuma bimaculata 
Phelsuma madagascariensis 
Ebenavia inunguis 
Zonosaurus aeneus 

Non-marine molluscs 

Macrochlamys stump fii 
Tropidophora tricarinata 
Ampelita xyster a 



Mantipus laevipes 
Platypelis pollicaris 



Mabuya gravenhorsli 
Amphiglossus melano pleura 
Chamaeleo nasutus 
Sanzinia madagascariensis 



Kalidos oleatus 
Helicophanta magnified 



NAME Reserve Naturelle Integrate de TSARATANANA (No.4). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927. 

GEOGRAPHICAL LOCATION 13°49'-14°05 , S, 48°44'-59'E. To the south-east of Ambanja 
in the province of Antseranana. 

ALTITUDE 700-2876 m (Mt. Maromokotra, the highest summit in Madagascar). 

AREA 48 622 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES Large mountain group of crystalline rocks and volcanic formations 
of the Miocene. Mont Maromokotra, the highest mountain in Madagascar, is found in this 
reserve. The climate has the same characteristics as the Malagasy rain forest and is extremely 
wet. During summer, especially the monsoon season (end of November to the beginning of 
May), there are virtually daily torrential rainstorms. From May to October, the summits are 
covered with fog accompanied by drizzle and fine rain. October and November are relatively 
dry. 

VEGETATION In general, the vegetation consists of primary and secondary tropical 

evergreen forests of both high and low altitude with both lichens and ericaceous species 
common. The flora is rich in endemics especially at high altitude. From 1000 to 2200 m, the 
principal plants found are: Podocdrpus mdddgdscdriensis, Cdtidrium, Aphloid theiformis, 
Rdvensdrd, Ocoted, Beilschmiedid oppositifolid, Mdlledstrum, Noronhid, Erythroxylum 
corybosum, Dichdetdntherd, EledCdrpus, Coffed tsaratananae, Gardenia, Peddiea involucrata, 
Buddleia, Senecio, Vernonia, Oncostemum, Panicum uvulatum, Poecilostachys tsaratananensis, 
Oplismenus, Leptaspis cochleata and various members of the families Acanthaceae, Labiaceae, 
especially Coleus, and Urticaceae, notably Piled. Tree ferns of the family Cyatheaceae do not 
seem to be very abundant; however this appears to be optimum habitat for epiphytes such 
as Peperomid, Kalanchoe, Medinilla, Viscum, Rhipsalis and numerous ferns and orchids. From 
2000 to 2200 m there is a belt of virtually pure giant bamboo. Above 2200 m there is moss 
forest with Araliaceae, Cunoniaceae (Weinmannia), Compositae, Ericaceae (Agauria, Philippia), 
Sterculiaceae (Dombeya), and Taxaceae (Podocarpus rostratus 

and P. madagascariensis). Chrysalidocarpus (Palmae) is also found. Ephiphytes are very 
common. Understorey, when it exists, is composed of small trees: Schismatoclada, 
Helichrysum, Philippia. Around 2600 m there is a second band of bamboos, then shrubby and 
herbaceous vegetation on the summits, secondary and depauperate through fire, of 

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An environmental profile of Madagascar 



blackened Philippia in 
scattered Helichrysum). 



a grassy lawn (Danthonia, Bromus, Anthoxanlhum, with 



DISTURBANCES OR DEFICIENCIES Some illicit cultivation; however, apart from the 
burned summits, the reserve is largely intact, as the very cold winter temperatures and the 
steepness of the terrain limit attempts at planting. Formerly, parts of the reserve were 
adversely affected by illicit plantations of tobacco and Indian hemp. In 1968 and 1969, a 
police operation was organised to destroy the plantations and arrest the miscreants. These 
people were sufficiently well organized to make entry to the reserve dangerous if unarmed and 
unaccompanied. Outside the plantations were found herds of zebu cattle belonging to the same 
people. It is not known if this has been a problem more recently. 

SCIENTIFIC RESEARCH Numerous collecting trips have been carried out. 

VISITOR AND SCIENTIFIC FACILITIES None. 

PRINCIPAL REFERENCE MATERIAL 

Albignac, R. (1970). Mammiferes et oiseaux du Massif de Tsaratanana. Mem. ORSTOM 

37: 223-229. 
Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 

Madagascar. In: Comptes rendus de la Conference internationale sur la Conservation de la 

Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 

IUCN, Switzerland. 
Milon, P. (1957). Etude d'une petite collection d'oiseaux du Tsaratanana. Naturaliste 

Malgache 3(2): 167-183. 

STAFF Two full-time forest agents. 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Headquarters are at Ambanja with a 
guard post at Mangindrano. 

FAUNA 



Birds 

Lophotibis cristata 
Buteo brachypterus 
Margaroperdix madagarensis 
Turnix nigricollis 
Dryolimnas cuvieri 
Sarothrura insularis 
Gallinago macrodactyla 
Streptopelia picturata (R) 
Alectroenas madagascariensis 
Coracopsis nigra (R) 
Coua reynaudii 
Coua caerulea 
Cuculus rochii (R) 
Caprimulgus ennaratus 
Alcedo vintsioides (R) 
Ipsidina madagascariensis 
Atelornis crossleyi 
Leptosomus discolor (R) 
Philepitta castanea 
Neodrepanis cor us cans 
Mirafra hova 
Phedina borbonica (R) 
Motacilla flavivenlris 
Coracina cinerea (R) 



Phyllastrephus zosterops 
Phyllastrephus cinereiceps 
Hypsi petes madagascariensis (R) 
Tylas eduardi 

Calicalicus madagascariensis 
Leptopterus viridis 
Copsychus albospecularis 
Monticola sharpei 
Neomixis viridis 
Hartertula flavoriridis 
Crossleyia xanthophrys 
Nesillas typica 
Dromaeocercus brunneus 
Dromaeocercus seebohmii 
Newtonia brunneicauda 
Pseudobias wardi 
Terpsiphone mutata (R) 
Nectarinia souimanga (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Lonchura nana (R) 
Ploceus nelicourvi 
Foudia madagascariensis (R) 
Dicrurus forficatus (R) 



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Protected areas 



Mammals 

Lemur macaco 
Lemur rubriventer 
Cheirogaleus major 
Lepilemur sp. 
Microgale sp. 

Amphibians 

*Mantipus guenther peter si 
Platypelis pollicaris 
Stumpffia psologlossa 
Mantidactylus elegans 

Reptiles 

Lygodactylus rarus 
*Chamaeleo tsaratananensis 



Hapalemur griseus 
Lemur fulvus 
Phaner furcifer 
Microgale talazaci 
Nesomys cf N. rufus 



*Platyhyla alticola 

*Platypelis tsaratananaensis 
Mantidactylus asper 
Mantidactylus granulatus 



Chamaeleo guibe 
*Amphiglossus tsaratananensis 



Nonmarine Molluscs 

This reserve is undoubtedly the most important area in Madagascar for molluscs. 



* Acroplychia culminans 
* ' Acroptychia pauliani 
*Acropytchia pauper 
*Tropidophora dingeoni 
*Tropidophora puerilis 

Tropidophora tricarinata 
*Tropidophora vuillemini 
*Clavator dingeoni 

Clavator moreleti 
*Clavator pauliani 
*Kalidos amicus 

Kalidos andapaensis 
*Ampelita caduca 
*Ampelita culminans 

Ampelita futura 

Ampelita gaud ens 

Ampelita lamarei 

Ampelita lanxava 

* Ampelita parva 

* Ampelita pauliani 
Ampelita perampla 

*Sitala acuta 
*Sitala culminis 



Kalidos cleamesi 
*Kalidos decaryi 

Kalidos humbloti 
*Kalidos merschardti 
*Kalidos milloti 
*Kalidos montis 

Kalidos oleatus 
*Kalidos secans 
*Kalidos tsaratananensis 
*Cyathopoma pauliani (Andamy) 

Macrochlamys stumpfii 
* Ampelita bathiei 
*Sitala delaportei 

Sitala elevata 
*Sitala roedereri 
^Microcystis madecassina 
*Microcystis nitelloides 
*Microcystis tangens 

Vitrina madagascariensis 

Edentulina metula 
*Pilula excavata 
*Pilula madecassina 



NAME Reserve Naturelle Integrate de L'ANDRINGITRA (No.5). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927; the present boundaries were fixed by Classement 
66-242, June 1st, 1966. 

GEOGRAPHICAL LOCATION 22°07'-2rS, 46°47 , -47 <, 02'E. South of Ambalava in the 
Province of Fianarantsoa. The village of Antanifotsy is situated on the edge of the reserve. 

ALTITUDE 1200-2658 m. 



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An environmental profile of Madagascar 

AREA 31 160 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The Andringitra mountain chain which constitutes the reserve is a 
very hilly, granitic massif in the form of a V with unequal length branches; where these meet 
the massif forms a plateau at about 2500 m with very uneven relief. The massif contain the 
sources of many streams and has the coldest climate of any of the Madagascan mountains, with 
severe frosts recorded at high altitude and several snowfalls at above 2500 m in the last 40 
years; lower altitudes in the reserve generally have the characteristic cool climate of the high 
plateaux with 3 or 4 dry months. Annual rainfall is 1500-2000 mm. On the eastern slopes, the 
climate is rather more equable. 

VEGETATION Several vegetation types occur within the reserve, varying principally with 
altitude, but also from east to west. The non-degraded vegetation is classified as follows: from 
700 to 800 m are areas of eastern lowland rain forest, characterised by Dalbergia baroni; from 
800-1600 m is found mid-altitude rain forest, with Eugenia, Tambourissa and Allocarpus; 
above this, from 1500-2000 m., is montane sclerophyll and lichen forest, characterised 
by Schefflera, Weinmannia and Brachylaena, and ericoid scrub with Philippia which can reach 
4-5 m in height. At these altitudes there are also scattered peaty depressions (notably carrying 
the remarkable Restio madagascariensis) and 'xerophytic lawn'. On the crags is a xerophytic 
flora with Aloe, Kalanchoe and Helichrysum. The whole area is rich in endemics: 80% of the 
flora of the humid depressions and the rocks is endemic to the massif. Much of the natural 
vegetation in the western part of the reserve has been destroyed by fire, but that which 
survives is characteristic of the western Madagascan domain. 

ZONING None. 

CONSERVATION MANAGEMENT The boundary is marked by 33 permanent marker stones, 
although these are not clearly visible; the reserve is patrolled on foot by three agents of the 
Direction des Eaux et Forets. There is a 30 km fire break along the western boundary of the 
reserve near the village of Ankazomby, although this has apparently only been partially 
effective. 

DISTURBANCES OR DEFICIENCIES Fire is the principal threat to the reserve; most 
burning within the reserve is the result of spread of uncontrolled fires started on pasture land 
outside the reserve, although some are deliberately started to cause damage and others may be 
caused by lightning. Around 500 ha of the summit zone was burnt in 1983 as was some 44 ha 
on the Andohariana Plateau; in 1982 around 1250 ha in the Antombohobe area was burnt. 
Cattle are present within the reserve and cause damage to the natural vegetation, particularly 
the rock-dwelling communities. The Riantahy and Rianvavy waterfalls, two areas reportedly 
of great beauty and historical interest, have been damaged by watercourse management under 
the Mamoly management project, designed to improve irrigation and rice production in the 
Antanifotsy village region. Management involves regulating the water flow at the Riantahy 
waterfall. To this end a dam and reinforced conduit have been built, and construction of a 
3 km long canal, scheduled to be completed in 1987, has begun; all this work has been carried 
out within the reserve. The agents of the Direction des Eaux et Forets lack camping equipment 
and transport and are thus prevented from effectively patrolling the reserve. Ring-tailed 
Lemurs Lemur catta are captured alive to sell to people passing through the area. 

SCIENTIFIC RESEARCH Study of highland ecosystems in 1970 for RCP 225 (Paulian et al., 
1971). 

VISITOR AND SCIENTIFIC FACILITIES Access is difficult: there is a 50 km track from 
Ambalavao to Antanifotsy which is in bad condition and is damaged each wet season and a 
40 km dry-season track to Ambaratra Antambohobe. There are several foot paths within the 
reserve, including one running 25 km from Antanifotsy to Pic Boby. 



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Protected areas 



PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 

Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 

Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 

IUCN, Switzerland. 
Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF 

programme in Madagascar. Unpd. report, 62 pp. 
Paulian, R., Betsch, J-M., Guillaumet, J-L., Blanc, C. and Griveaud, 

225. Etude des ecosystems montagnards dans la region malgache. 

l'Andringitra. 1970-1971. Geomorphologie, climatologie 

vegeteaux. Bulletin de la Societe d'Ecologie II (2-3): 189-266. 



Protected areas 

P. (1971). RCP 
I. Le massif de 
et groupements 



STAFF 1 agent and 2 full-time auxiliaries. 
BUDGET Salaries paid by the government. 



LOCAL PARK OR RESERVE ADMINISTRATION 

guardpost at Ivohibe. 



Headquarters at Ambalavo and a 



FAUNA 



Birds 

Tachybaptus pelzelnii 
Anas melleri 
Polyboroides radiatus 
Buteo brachypterus 
Falco newtoni (R) 
Sarothrura insularis 
Streptopelia picturata (R) 
Cuculus rochii (R) 
Leptosomus discolor (R) 
Atelornis pitt ides 
Phedina borbonica (R) 
Motacilla flaviventris 



Hypsi petes madagascariensis (R) 
Copsychus albospecularis 
Monticola sharpei 
Acrocephalus newtoni 
Nesillas typica 
Cisticola cherina 
Dromaeocercus brunneus 
Neomixis tenella 
Nectarinia souimanga (R) 
Zosterops maderaspatana 
Foudia madagascariensis (R) 



Mammals 

Microcebus rufus 
Avahi laniger 
Lepilemur microdon 
Lemur catta 
Lemur fulvus 
Varecia variegata 
Cryptoprocta ferox 
Tenrec ecaudatus 



Setifer setosus 
Microgale dobsoni 
Microgale drouhardi 
Leptogale gracilis 
Oryzorictes tetradactylus 
Brachyuromys betsiloensis 
Brachyuromys ramirohitra 
Eliurus myoxinus 



Amphibians 

*Anodonthyla montana 
Mantipus inguinalis 
Plethodontohyla tuberata 
Mantidactylus aerumnalis 
Mantidactylus argenteus 

*Mantidactylus blanci 
Mantidactylus decaryi 

Mantidactylus elegans 
* Mantidactylus madecassus 

Mantidactylus redimitus 
*Boophis brygooi 

Boophis microtympanum 



*Mantipus bipunctatus 
Plethodontohyla notosticta 
Pseudohemisus madagascariensis 
Mantidactylus aglavei 
Mantidactylus asper 

*Mantidactylus bourgati 
Mantidactylus domerguei 

Mantidactylus lugubris 
Mantidactylus microtympanum 
Mantidactylus tricinctus 
*Boophis laurenti 



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An environmental profile of Madagascar 

Reptiles 

Leioheterodon madagascariensis 

Nonmarine Molluscs 

Ampelita covani *Tachyphasis milloti 

Ampelita petiti Macrochlamys stump fii 

*Imerinia fischeri Helicophanta gloriosa (foret d'Fivanona) 

Amphipoda: 

Austroniphargus bryophilus (Pic Boby) 



NAME Reserve Naturelle Integrate de LOKOBE (No.6). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927, though the Foret de Lokobe was constituted as a 
reserve as early as 1913. 

GEOGRAPHICAL LOCATION On the south-eastern side of the island of Nosy Be, with the 
coast forming the southern edge of the reserve. 13°23'-25'S, 48°18'-20'E. 

ALTITUDE 0-430 m (the highest point on the island). 

AREA 740 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The island of Nosy Be is formed of a neogenous basaltic block and 
marine sediments of the upper Lias. The relief is fairly tortuous in places. The reserve has an 
important role in the local water network. Climate is characteristic of the 'Sambirano' Domain, 
with a relatively well-marked dry season of 3 to 4 months, lower annual rainfall than on the 
east coast and fairly high temperatures (mean of over 15°C in the coldest month). 

VEGETATION The reserve contains much of the remaining forest on the island. This is a 
dense, humid forest characterised by species of the families Chlaenaceae (endemic to 
Madagascar), Myristicaceae and the genus Anthostema. Biologically and physionomically, it 
resembles the eastern rain forest but is distinguished by the presence of numerous endemics. 

ZONING The reserve boundaries are clearly delimited, and obviously marked. 

CONSERVATION MANAGEMENT There are proposals to increase the effectiveness of the 
reserve both by increasing its size, and by providing vehicles for the staff. Use of surrounding 
areas for tourism could be a valuable source of income. 

DISTURBANCES OR DEFICIENCIES The reserve is very vulnerable because of its small 
size. The proposed buffer zone, which is not formally protected, is being cut down for 
cultivation of rice and manioc. In 1972 it was reported that poaching of lemurs was a problem, 
incursions being made into the reserve by boat. 

SCIENTIFIC RESEARCH Numerous collecting expeditions have been carried out. 

VISITOR AND SCIENTIFIC FACILITIES The Centre National de Recherches 

Oceanographiques owns a laboratory close to the reserve, but the work has always been 
orientated towards the marine fauna. 



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Protected areas 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

STAFF Only one agent. 

BUDGET Salaries paid by the government. In 1982, a WWF Tropical Forest Campaign grant 
of US$ 22 000 over 2 years was allocated to buy essential equipment and assist in the protection 
of 150-200 ha of the planned buffer zone in the northern part of the reserve. 

LOCAL PARK OR RESERVE ADMINISTRATION At Hell-ville on Nosy Be. 

FAUNA 

Birds 

The following species have been recorded by O. Langrand (in litt., 28.10.86). 



Aviceda madagascariensis 
Haliaeetus vociferoides 
Polyboroides radiatus 
Accipiter francesii (R) 
Buteo brachypterus 
Falco newtoni (R) 
Margaroperdix madagarensis 
Turnix nigricollis 
Streplopelia picturata (R) 
Treron austral is (R) 
Alectroenas madagascariensis 
Coracopsis nigra (R) 
Agapornis cana 
Cuculus rochii 
Coua cristata 
Centropus toulou (R) 
Otus rutilus (R) 

Caprimulgus madagascariensis (R) 
Zoonavena grandidieri (R) 
Alcedo vintsioides (R) 
Eurystomus glaucurus (R) 

Mammals 

Lepilemur dorsalis 
Microcebus rufus 



Leptosomus discolor (R) 
Motacilla flaviventris 
Coracina cinerea (R) 
Phyllastrephus madagascariensis 
Hypsi petes madagascariensis (R) 
Calicalicus madagascariensis 
Vanga curvirostris 
Leptopterus viridis 
Leptopterus chabert 
Leptopterus madagascarinus (R) 
Copsychus albospecularis 
Nesillas typica 
Newtonia brunneicauda 
Neomixis tenella 
Terpsiphone mutata (R) 
Nectar inia soui manga (R) 
Nectarinia notata (R) 
Zoslerops maderaspatana 
Lonchura nana (R) 
Foudia madagascariensis (R) 
Dicrurus forficatus (R) 



Lemur macaco 
Microcebus murinus 



Amphibians 

The following taxa are listed for Nosy Be in general, no data are available on the 
herpetofauna of Lokobe in particular. Platypelis milloti is an exception; it has been 
recorded only in the Lokobe Reserve. 



Cophyla phyllodactylus 
Rhombophryne testudo 
Laurentomantis horrida 
Boophis madagascariensis 



*Platypelis milloti 
Stumffia psologlossa 
Mantidactylus granulatus 

'Hyperolius' nossibeensis 



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An environmental profile of Madagascar 



Reptiles 

The following taxa are listed for Nosy 
herpetofauna of Lokobe in particular. 

Ebenavia inunguis 
*Lygodactylus heterurus 

Phelsuma laticauda 
*Paroedura oviceps 

Uroplatus fimbriatus 

Brookesia ebenaui 
*Brookesia minima 

Brookesia superciliaris 

Chamaeleo parsoni 
*Amphiglossus stumpffi 

Zonosaurus boettgeri 

Zonosaurus rufipes 
*Typhlops madagascariensis 

Acrantophis madagascariensis 

Langaha nasuta 

Lycodryas arctifasciatus 

Micropisthodon ochraceus 

Liopholidophis stumpffi 



Be in general, no data are available on the 



Geckolepis maculata . 
Phelsuma dubia 
Phelsuma madagascariensis 
Paroedura stumpffi 
Uroplatus ebenaui 

*Brookesia legendrei 
Brookesia stumpffia 
Chamaeleo boettgeri 
Amphiglossus polleni 
Paraconlias hildebrandti 
Zonosaurus madagascariensis 
Ramphotyphlops braminus 

*Typhlops reuteri 
Ithycyphus miniatus 
Liophidium rhodogaster 
Lycodryas gaimardi 
Pararhadinea melanogaster 



Nonmarine Molluscs 

Some of these species are recorded from 'Nosy Be' and may not occur in Lokobe. 



Tropidophora aspera 

Tropidophora cuvieriana 

Tropidophora deshayesiana 
*Tropidophora felicis 

Tropidophora fuscula (Lokobe) 

Tropidophora ligata non endemic 

Tropidophora milloti 

Tropidophora tricarinata 

Tropidophora vittata 

Clavator moreleti 

Helicophanta amphibulima (Lokobe) 

Helicophanta oviformis (Lokobe) 

Kalidos lamyi (Lokobe) 
*Sitala brancsiki (Lokobe) 
*Sitala filomarginata (Lokobe) 

Edentulina stumpfii 

Macrochlamys stumpfii 

Acmella parvula 

Cleopatra colbeaui 



Truncatella guerini 
Gastrocopta seignaciana 
Nesopupa minutalis 
Cecilioides mariei 
Ampelita galactostoma 
Ampelita gaud ens 
Ampelita omphalodes (Lokobe) 
* Ampelita stumpfii (Lokobe) 
Desmocaulis subaspera 
Drepanocaulis plateia 
Drepanocaulis tetragonalis 
Euconulus micra 
Opeas soulaianus 
Elisolimax bella 
Imerinia grandidieri 
Imerinia sulfurea 
Imerinia verrucosa 
Clithon spiniperda 



NAME Reserve Naturelle Integrate de L'ANKARAFANTSIKA (No.7) 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED The integral reserve was established by a decree of 31 December 
1927, and updated by decree 66-242 of 30 June 1966. 

GEOGRAPHICAL LOCATION In Mahajanga Province 40 km north-west of Ambato-Boeni. 
15°59'-16°22'S, 45"56'-47°12'E. 



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Protected areas 

ALTITUDE 75-390 m. 

AREA 60 520 ha, bordering the Ampijoroa Forest Station to the west which covers ca 
20 000 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES Very rugged relief to the east and south, forming cliffs in places. 
Towards the west and north, the plateau descends gently. The soils of the plateau are very 
sandy. Precipitation is between 1000 and 1500 mm a year with a marked dry season of 5-6 
months (May-November). Mean annual temperature is ca 26°C, with maximum 35°C and 
minimum 17°C. The reserve protects a sample of habitats typical of the arenaceous soils of 
western Madagascar, and also protects the catchment of one of Madagascar's most important 
rice growing areas. Problems occur in the paddy fields downstream when sand eroded from 
cleared areas is carried by the rivers. 

VEGETATION Still largely covered in the original forest vegetation. Forest is dense and dry 
of the series Dalbergia-Commiphora-Hildegardia. Numerous Leguminosae and Myrtaceae. 
Some species adapted to dry environments such as Pachypodium, and members of the families 
Ampelidaceae and Passifloraceae. Numerous lianes, but epiphytes are virtually absent. The 
forest is deciduous, and contains a wide variety of trees and shrubs at a high density (about 170 
species of 35 families). 

ZONING None, althuogh the reserve is bordered by six buffer zone areas. 

CONSERVATION MANAGEMENT The reserve itself is not managed at present. The 
forestry station has a nursery and plant growth trials area; there is a good network of trails 
through the forest station and part of the area is used for research projects. 

DISTURBANCES OR DEFICIENCIES The reserve and forest station are being encroached 
by clearance to create pastures, by charcoal burning, and, to a lesser extent, by clearance for 
crops in river valleys and slopes on the Antananarivo to Mahajanga road. The buffer zone 
areas have suffered forest loss and the forest edges are being pushed back into the reserve, 
particularly in the north and east. Fire affected the western quarter of the reserve in 1983. 
Poaching appears to be low-level at present (1986) although may become more of a problem in 
the future. There are feral cattle within both the reserve and the forest station. The reserve is 
undermanned. 

SCIENTIFIC RESEARCH Some primate studies have been made in the reserve, particularly 
on range sizes and diet; the University of Madagascar also has study areas here. Entomological 
research has been carried out in the area. 

VISITOR AND SCIENTIFIC FACILITIES The forestry station at Ampijoroa is used by 
research workers in the area. 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

IUCN/WWF Project 1911. Protection and development of the Ankarafantsika Nature Reserve. 

Martin, C. (1982). Rapport de la mission technique WWF/IUCN a Madagascar 1981. 
IUCN/WWF, Gland, (contains a list of birds and mammals found within the reserve). 

Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF - Protected areas 
programme in Madagascar. Unpd. report, 62 pp. 

STAFF Two: a Chef de Station at Ampijoroa and a full-time auxiliary. 

BUDGET Salaries paid by the government. WWF has provided funds for reserve management 
since 1980. 

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An environmental profile of Madagascar 

LOCAL PARK OR RESERVE ADMINISTRATION The headquarters is at Bevazaha. 

FAUNA 



N.B. Some of the species listed 
immediately adjacent to the R.N.I. 



below have been recorded from Ampijoroa which lies 



Birds 

Tachybaptus pelzelnii 

Ardeola idae (R) 

Lophotibis cristata 

Anas melleri 

A viced a madagascariensis 

Haliaeetus vociferoides 

Polyboroides radialus 

Accipiter madagascariensis 

Accipiter francesii (R) 

Buteo brachypterus 

Falco newtoni (R) 

Margaroperdrix madagarensis 

Mesitornis variegala 

Turnix nigricollis 

Dryolimnas cuvieri (R) 

Plerocles personatus 

Slreptopelia picturata (R) 

Treron australis 

Agapornis cana 

Coracopsis nigra (R) 

Coracopsis vasa (R) 

Coua gigas 

Coua coquereli 

Coua ruficeps 

Coua cristata 

Cuculus rochii (R) 

Centropus loulou (R) 

Olus rutilus (R) 

Asio madagascariensis 

Caprimulgus madagascariensis (R) 

Zoonavena grandidieri (R) 

Alcedo vintsioides (R) 

Mammals 

Avahi laniger 
Lepilemur edwardsi 
Cheirogaleus medius 
Lemur fulvus 
Macrotarsomys ingens 
Tenrec ecaudatus 
Pteropus rufus 



Ipsidina madagascariensis 
Eurystomus glaucurus (R) 
Leptosomus discolor (R) 
Philepitta castanea 
Mirafra hova 
Phedina borbonica (R) 
Molacilla flaviventris 
Coracina cinerea (R) 
Phyllastrephus madagascariensis 
Hypsi petes madagascariensis 
Calicalicus madagascariensis 
Schetba rufa 
Vanga curvirostris 
Xenopirostris damii 
Falculea palliata 
Leptopterus chabert 
Leptopterus viridis 
Leptopterus madagascarinus (R) 
Copsychus albospecularis 
Acrocephalus newtoni 
Nesillas typica 
Cisticola cherina 
Newtonia brunneicauda 
Neomixis tenella 
Terpsiphone mutata (R) 
Nectarinia notata (R) 
Nectarinia souimanga (R) 
Zosterops maderaspatana 
Lonchura nana ( R) 
Ploceus sakalava 
Foudia madagascariensis (R) 
Dicrurus forficatus (R) 



Lemur mongoz 
Microcebus murinus 
Propithecus verreauxi 
Cryptoprocta ferox 
Macrotarsomys bastardi 
Setifer setosus 



Quoted records of Lepilemur ruficaudatus should almost certainly be ascribed to L. edwardsi 

Pseudohemisus granulosus 



Amphibians 

Dyscophus insularis 
Mantidactylus wittei 



Reptiles 

*Brookesia decaryi 

*Pygomeles petteri 

Phelsuma dubia 



*Chamaeleo angeli 
Langaha nasuta 



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Protected areas 



Nonmarine Molluscs 

Ampelita omphalodes Helicophanta amphibulima (Bevahara) 

Ampelita pfeifferi Helicophanta oviformis (Bevaraha) 

Euconulus micra Acroptychia millotti 

Edentulina gaillardi 

Nonmarine Crustacea 
Isopoda: 

Armadillo silvivagans (Tsaramandroso) 

Lepidoptera 
Papilionidae 

Papilio morondavana Papilio grosesmithi 



NAME Reserve Naturelle Integrate du TSINGY DE NAMOROKA (No.8). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927. 

GEOGRAPHICAL LOCATION To the south of Soalala in the province of Mahajanga. 
16°19'-30'S, 45°16'-25 , E. 

ALTITUDE 180-370 m. 

AREA 21 742 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The area comprises a calcareous massif (karst) with many cliffs and 
numerous caves and springs. Average rainfall is between 1000 and 1500 mm a year, with a 
distinct dry season between May and November. Mean temperature in the coolest month is 
above 20°C. 

VEGETATION Like the Tsingy de Bemaraha (R.N.I. No.9), the reserve is a mosaic of dense 
dry forest, savanna and vegetation adapted to the calcareous karsts, belonging to 
the Dalbergia-Commiphora-Hildegardia association. The mean height of the trees is 
12-15 m. Adansonia rubrostipa is especially frequent. Many xerophytic and crassulacean 
plants. The area has a spring-fed stream with a remarkable aquatic flora. This reserve has 
lower species diversity than Tsingy de Bemaraha, whilst containing essentially the same 
biotopes. 

ZONING None. 

CONSERVATION MANAGEMENT Some 14 km of footpaths are delimited within the reserve 
as passable at all times of the year. 

DISTURBANCES OR DEFICIENCIES Fires are frequent during the dry season. The human 
population around the reserve is fairly low and there is only one important village - 
Vilanandro. In 1972 it was noted that the local inhabitants were largely indifferent to the laws 
protecting the reserve, which was inadequately guarded; no fady (local taboo) protected the 
animals or plants of the region. Plantations of indian hemp have been found in the interior of 
the reserve and zebu cattle also occur within the reserve. 

SCIENTIFIC RESEARCH No research appears to have been carried out recently. 

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An environmental profile of Madagascar 
VISITOR AND SCIENTIFIC FACILITIES None. 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference international sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

STAFF One agent and a full-time auxiliary. 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Reserve headquarters at Vilanandro. 

FAUNA 



Birds 

Dryolimnas cuvieri 

Treron australis (R) 

Alectroenas madagascariensis 

Agapornis cana 

Coua ruficeps 

Coua coquereli 

Otus rutilus 

Coua cristata 

Coua gigas 

Otus rutilus (R) 

Mammals 

Lepilemur edwardsi 
Propithecus verreauxi 

Reptiles 

*Brookesia bonsi 

Nonmarine Molluscs 

Acroptychia grandidieri 
Kalidos aequivocus 
Kalidos bournei 
Helicophanta oviformis 
*Bathia madagascariensis 



Philepitta schlegeli 
Motacilla flaviventris 
Schetba rufa 
Vanga curvirostris 
Copsychus albospecularis 
Neomixis tenella 
Nesillas typica 
Zosterops maderaspatana 
Ploceus sakalava 



Microcebus murinus 
Lemur fulvus 



Georissa aurata 
Boucardicus petiti 
Tropidophora semidecussata 
Ampelita namerokensis 



NAME Reserve Naturelle Integrate du TSINGY DE BEMARAHA (No.9). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927. 

GEOGRAPHICAL LOCATION To the east of Antsalova in the region of Antsingy, 

18°13'-19°07 , S, 44°34'-57'E. 

ALTITUDE 75-700 m. 

AREA 152 000 ha, the largest natural reserve in Madagascar. 



132- 



Protected areas 

LAND TENURE Government land. 

PHYSICAL FEATURES The reserve forms a part of the Antsingy region, a limestone karst 
area of very dissected relief with many caves and springs. Much of the eastern edge of the 
region is delimited by the Bemaraha cliff, several tens of kilometres long and 300 to 400 m 
high. To the east of the reserve there are three north-south flowing rivers separated by 
successive ridges, while the whole western region of the reserve forms a plateau with rounded 
hillocks which slopes away, with decreasing steepness, to the west. The climate is generally 
dry and there are 7 or 8 dry months. Temperatures decrease from north to south but are 
always above 20°C. 

VEGETATION Vegetation is characteristic of the calcareous karst regions of the west, with 
many species unique to this formation, such as Diospyros perrieri (the ebony of the west 
coast), Delonix regia, and other species of the genus Delonix, and Musa perrieri (the only wild 
banana in Madagascar). Also baobabs Adansonia, and xerophytic plants such as Aloe. Other 
notable families include Flacourtiaceae, Orchidaceae, Leguminosae, Euphorbiaceae, 
Annonaceae, Bombacaceae and Moraceae. Climate and vegetation are very similar to that of 
the R.N.I, du Tsingy de Namoroka (No. 8), and the main vegetation types are dense, dry forest 
and savannah, but the much larger area and the greater height of the karst relief make the 
vegetation richer here. 

ZONING None. 

CONSERVATION MANAGEMENT There appears to be no active conservation management 
at present. 

DISTURBANCES OR DEFICIENCIES Access to much of the reserve is very difficult, 
resulting in many areas being naturally protected. However, a track crosses the reserve from 
east to west and accessible valleys are populated with zebu cattle; there are also reportedly 
illegal settlements within the reserve. Poaching has occurred, though it is not known if this 
continues to be a problem. 

SCIENTIFIC RESEARCH Very little. The area was apparently surveyed in the 1930s, 
though it is not known if the results have been published; in 1972 it was reported that a recent 
evaluation of the vegetation had been carried out, though further details are lacking. The 
reserve is of considerable archaeological interest, containing numerous ancient cemeteries. 

VISITOR AND SCIENTIFIC FACILITIES None. 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

Leandri, J. (1938). La foret d'Antsingy. La Terre et la Vie: 18-27. 

STAFF One agent and two full-time auxiliaries. 

BUDGET Salaries provided by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Reserve headquarters is at Antsalova with 
a guard post at Bekapaka. 

FAUNA 

Birds 

Coua ruficeps Lophotibis cristata 

Coua gigas Vanga curvirostris 

PTachybaptus rufolavatus (record questioned) 

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An environmental profile of Madagascar 

Mammals 

Lepilemur edwardsi 
Microcebus coquereli 
Propithecus verreauxi 
Phaner furcifer 

Reptiles 

*Brookesia perarmata 



Microcebus murinus 
Lemur fulvus 
Hapalemur griseus 



Lygodaclylus klemmeri 



Nonmarine Molluscs 

The Gorges and grottes de Salapango are in this area, but probably not actually in the 
reserve. The following species occur in the Bemaraha region and may also occur in the 
gorges de Salapango (see below): 



Tropidophora bemaraensis 
Tropidophora chavani (Salapango) 
Tropidophora filopura " 
Tropidophora peliti 
Tropidophora vignali " 



Boucardicus petiti (near R. Nameroko) 
Edentulina battistini 
Kalidos bournei 
Tropidophora semidecussata 
Tropidophora pyrostoma 
(near Miandrivazo) 



Gorges (and grottes) de Salapango - these species are specifically recorded from this locality: 



Tropidophora chavani 
Tropidophora filopura 
*Tropidophora petiti 
Tropidophora vignali 
Edentulina battistini 
Edentulina stumpfii 



*Ampelita milloti 
Ampelita namerokoensis 
Chondrocyclus mamillaris 
Acroptychia bathiei 
Kalidos bournei 
Helicophanta oviformis 



NAME Reserve Naturelle Integrate de TSIMANAMPETSOTSA (No. 10). 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 31 December 1927. 

GEOGRAPHICAL LOCATION 24°02'-irS, 43°36'-5rE. 100 km south of Toliara in the 
province of Toliara. 

ALTITUDE 10-160 m. 

AREA 43 200 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The western part of the reserve comprises the shallow, brackish 
Tsimanampetsotsa lake (20 x 3 km), saturated with sulphates of lime. The lake, which has 
shores of virtually unvegetated arenaceous soil, is aligned north-south and lies about 7 km from 
the west coast. To the east of the lake is xerophytic forest on calcareous bedrock. There are 
numerous underground caves. The climate is dry with precipitation below 400 mm a year and 
9-11 dry months. Amount and timing of rainfall can be very variable. Other forms of 
precipitation are likely to have a significant impact on the vegetation. The minimum 
temperature in the coolest months is between 15° and 20°C. 



■134- 



Protected areas 

VEGETATION Xerophtyic scrub on the calcareous plateau, and typical south-western brush 
formation on the arenaceous soils. The vegetation is a remarkable assembly of Didiereaceae (an 
endemic family of trees or shrubs with the habit of cacti or cactiform euphorbias) and 
Euphorbiaceae. Tree species 10-12 m high dominate a stratum of impenetrable scrub, rich in 
lianas. The ground stratum is sparse. The reserve covers a part of the very restricted 
distribution of Alluaudia montagnacii. Plants show a wide variety of adaptations to xerophytic 
conditions. Numerous species of Leguminosae, Combretaceae, Tiliaceae and Liliaceae are 
found. 

ZONING None. 

CONSERVATION MANAGEMENT The boundaries are not marked on the ground, though 
the reserve was apparently, at least up to 1972, still well respected, as a local 'fady' (taboo) 
acted on local villagers in conjunction with the official protection, and the villagers did not 
like accompanying visitors into the reserve. The particular characteristics of the site 
(calcareous plateau surrounded by very rocky hillsides covered with an uninflammable 
xerophytic bush and lakesides of practically bare soil) also shelter it from bush fires. The 
reserve is reportedly the only one without feral zebu cattle as there is apparently no, or very 
little, fresh water. There being no fish in the lake, there is no human disturbance for fishing. 
Overall, the reserve is therefore reported to be little disturbed and in good condition. The area 
in which it is situated is one of very low population density, and efficient surveillance could 
easily be carried out by controlling the exit to Androka and that to Toliara. The plateau whi-h 
borders the lake should be included in the reserve and it would be advisable, for example, to 
protect the potholes facing Itampolo in which lives Typhleotris madagascariensis, a legally 
protected blind fish. 

DISTURBANCES OR DEFICIENCIES Apparently relatively few (see above). 

SCIENTIFIC RESEARCH Little recent research appears to have been carried out. 

VISITOR AND SCIENTIFIC FACILITIES None. 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

Germain, L. (1935). Etude du reserve naturelle du lac Manampetsa [sic]. Ann. Sci. Nat. 
Zool. XVII: 421-481. 

Milon, P. (1950). Deux jours au lac Tsimanampetsoa [sic]. Observations ornithologiques. 
Naturaliste Malgache 2(1): 61-67. 

STAFF One full-time auxiliary. 

BUDGET None. 

LOCAL PARK OR RESERVE ADMINISTRATION None. 

FAUNA 

Birds 

Accipiter madagascariensis Agapornis cana 

Buteo brachypterus Cuculus rochii (R) 

Falco newtoni (R) Coua cursor 

Turnix nigricollis Coua cristata 

Charadrius thoracicus Coua ruficeps 

Pterocles personatus Coua verreauxi 

Streptopelia picturata (R) Centropus toulou (R) 

Coracopsis vasa (R) Otus rutilus (R) 

Coracopsis nigra (R) Ninox super ciliaris 

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An environmental profile of Madagascar 



Caprimulgus madagascariensis (R) 
Alcedo vintsioides (R) 
Leptosomus discolor (R) 
Mirafra nova 
Phedina borbonica (R) 
Coracina cinerea (R) 
Vanga curvirostris 
Xenopirostris xenopirostris 
Falculea palliata 
Leptopterus viridis 
Leptopterus chabert 
Monticola imerina 
Neomixis tenella 
Neomixis striatigula 



Nesillas lypica 
Thamnornis chloropetoides 
Cisticola cherina 
Newtonia amphichroa 
Newtonia brunneicauda 
Newtonia archboldi 
Terpsi phone mutata (R) 
Nectar inia souimanga (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Foudia madagascariensis (R) 
Lonchura nana (R) 
Ploceus sakalava 
Dicrurus forficata (R) 



Mammals 

Lepilemur leucopus (probably) 
Propithecus verreauxi 



Lemur catta 



Microgale pusilla has been found in owl pellets in the region (M. Nicoll, in litt., 28.10.86). 



Reptiles 

IPyxis arachnoides 



Geochelone radiata 



Nonmarine Molluscs 

Many of the following species were collected at lac 'Manampetsa': 



*Microcystis bathiei 
Kalidos lapillus 
Kalidos chastelli 
Clavator grandidieri 
Subulina manampetsaensis 



Leucotaenius favannii 
Leucotaenius procteri 
Georissa petiti 
Tropidophora philippiana 
Tropidophora semidecussata (fossil) 



The following species were recorded in the reserve during a survey in the 1930s (Petit, 1935; 
see also Part V.9. and Appendix 2). Names of some of these species have since been changed. 



Clavator favannei 
non endemic 
Subulina octona 
Subulina ferriezi 
Planorbis trivialis 
Planorbis crassilabrum 
Ligatella philippi 



Segmentina angusta 
Melania tuberculata 
Georissa petiti 
Truncatella teres 



Nonmarine Crustacea 
Amphipoda: 

Grandidierella mahafalensis 



Orchestia ancheidos 



An expedition carried out in the 1930s, as part of a series to inventory the reserves, recorded 
the following crustaceans (Monod, 1935): 

Ostracoda: 

Acoccypris capillata (not endemic) 
Tanaid?: 

Apseudes thaumastocheles (n.sp., blind, in chalky mud, ?endemic) 
Isopoda: 

Aphiloscia annulicornis (not endemic) 

Pyrgoniscus petiti (n.sp.) 
Amphipoda: 

Grandidierella megnae (not endemic) 



136- 



Protected areas 

To the north of this reserve is found the cave of Mitoho. This is an extremely important 
cave faunistically, and in 1935 it was reported that some measures had been taken for its 
conservation (Germain, 1935). It contains a remarkable collection of cave species of marine 
origin, as well as diurnal soft water species. The cave is on the edge of an ancient cliff near 
the plateau of Mahafaly and is connected with the water table circulating under the plateau 
(Paulian & Delamare Deboutteville, 1956). Since the cave is the only source of fresh water 
in the area, its fauna may easily become threatened (Paulian. 1983). The following endemic 
crustaceans are found there: 

Isopoda: 

Anopsilana poissoni 
Decapoda: 

Typhlopatsa pauliani 

Fish 

The blind cave fish, Typhleotris madagascariensis, is found in the limestone potholes near 
Itampolo; it has been recommended that the reserve be extended to cover this area. 



NAME Reserve Naturelle Integrate d'ANDOHAHELA (No.ll) 

MANAGEMENT CATEGORY I (Strict Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 11 June 1939. The area of the reserve was increased from 30 000 ha 
on 1 June 1966 by Decree 66-242. 

GEOGRAPHICAL LOCATION 24°30'-58 , S, 46°32'-52'E. 40 km north-west of Taolanaro in 
the extreme south of the country. 

ALTITUDE 100-1956 m 

AREA 76 020 ha, in three noncontiguous blocks, one (Parcel 1) 63 100 ha, one (Parcel 2) 
12 420 ha and the third (Parcel 3) 500 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES Parcel 1 varies in altitude from 100 m to 1956 m (Pic d'Andohahela); 
Parcel 2 from ca 110 m to 1005 m (Pic de Vohidagoro); Parcel 3 is at ca 125 m. Climate in 
parcel 1 is humid, with rainfall of 1500-2000 mm, no dry season and mean annual temperature 
of about 23°C; that in parcel 2 is much drier, with rainfall usually lower than 500 mm per 
annum and a dry season of 5-6 months. Parcel 1 is an important watershed, containing the 
source of over ten rivers, including the Mananara, rising at Anpamosira and flowing westward 
and the Manampanihy flowing east from Vohibe (O'Connor et al., 1985). The Mananara, 
which flows along the northern boundaries of Parcel 2, is the only permanent water source for 
that part of the reserve. 

VEGETATION Each of the three parcels has a distinct vegetation type. 

Vegetation in Parcel 1 is typical of submontane tropical rainforest, of which it constitutes the 
southernmost extension in Madagascar. Buttressed trees of up to 35 m occur, though generally 
tree height does not exceed 25 m. Genera characteristic of this forest type 
include Tambourissa, Symphonia and Dalbergia, with members of the families Lauraceae, 
Compositaceae and Rubiaceae represented on the higher slopes. The endemic family 
Humbertaceae is found within the reserve. Orchidaceae and Cycathaceae are common, and the 
epiphytic cactus Rhipsalis occurs. Epiphytes in general are abundant, and at higher altitudes 
mosses and lichens are found. 

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An environmental profile of Madagascar 

Parcel 2 consists mainly of spiny thorn forest with some bush and scrub and also some gallery 
forest along the Menanara River in the northern part of the reserve. The highest hills have no 
forest cover and are generally covered with tussock grass and other herbaceous vegetation, 
with Aloe and Pachypodium spp. In the thorn forest, the endemic genera Alluaudia 
and Didierea are well represented; one species of the former (A. ascendens) is endemic to the 
Mandrare region, as is the baobab Adansonia za (Bombaceae). Species of Euphorbiaceae, 
Leguminosae and Crassulaceae are also abundant. 

Parcel 3 has a high density of the endemic palm Neodypsis decaryi and was originally set up 
specifically to protect this species. It also has a belt of vegetation transitional between the 
spiny forest and the eastern rain forest. Leguminosae, particularly Acacia spp., are well 
represented as are Cucurbitaceae and Euphorbiaceae. There is some deciduous forest 
with Tamarindus indica along one of the non-permanent rivers, the Andehamara, and 
introduced Eucalyptus has become established along the eastern end of the parcel. 

ZONING None apart from the three defined areas. 

CONSERVATION MANAGEMENT Principal management at present is in the north-east of 
Parcel 1, near the village of Vohibaka, where a scheme to prevent fires spreading into the 
reserve has been started by the Chef de Cantonnement Forestier. Villagers have been using 
back fires to clear a 20 m wide firebreak which also deliniates the boundary of the reserve. 
Payment for this work up to the present has been in the form of food provided personally by 
the Chef de Cantonnement Forestier. 

O'Connor et al. have made several suggestions for improving management of the reserve: 1. 
Review of the present boundaries; there are large areas of the reserve which are devoid of 
forest and it is suggested that these be excluded from the reserve in exchange for equal areas of 
forest. This is particularly advocated for the thorn forest, of which there are considerable 
intact areas around parcel 2, as it is perceived as under increasing pressure for charcoal 
production and exploitation of Alluaudia wood. The boundaries of parcel 1 are in question at 
present as the original markers were incorrectly placed. 2. Clear delineation of boundaries, 
involving extension of the firebreak scheme around Vohibaka. The possibility of fencing 
parcel 3 should be investigated - its small size makes fencing feasible and its proximity to the 
main road makes it desirable. 3. An increase in the number of guards to ten, preferably 
employing local villagers, and provision of uniforms. 4. Provision of transport for the Chef de 
Cantonnement Forestier. 5. Promotion of local agricultural development schemes to alleviate 
pressure on the reserve. The critical need for funding for these activities is stressed. 

DISTURBANCES OR DEFICIENCIES All three parcels have deforested areas; in parcel 1 
these are presumed to be largely the result of bush fires, in parcel 2 a product of wood-cutting 
and in parcel 3 a combination of the two. The lower, flat areas are used to cultivate rice while 
the higher areas are used as cattle pasture, and are burned annually to provide new growth for 
grazing. In parcels 1 and 2 these areas are extensive, though few are of recent origin; in parcel 
1, however, bush fires are likely to be a problem near border villages, particularly near 
Eminiminy on the eastern side of the reserve. Slopes within the reserve here were noted by 
O'Connor et al. (1985) to be devoid of forest and it was thought likely that the villagers no 
longer considered them part of the reserve. There was also some evidence that the extent of 
cultivated land in the reserve was expanding in the southern end of parcel 1, north of the 
village of Isaka Ivondro; houses were noted within the reserve and new clearings were observed 
along the Ambahibe River. Fires have been recorded in the vicinity of Parcel 3 and could pose 
a serious threat to the area if not controlled. Livestock habitually graze within the reserve 
boundaries and wood-cutting is a problem where villages occur near to forested parts of the 
reserve. Wood taken from Parcel 1 is used mainly for fuel in homes whereas that taken from 
Parcel 2 is used for fuel and house building and is also taken for sale to markets in 
Ambovombe, Amboasary and Taolanaro. There is, however, as yet no evidence for large-scale 
logging within the reserve. Hunting is generally not a major problem, though may be of some 
importance in the north and east of Parcel 1 where cattle thieves hide and in parcel 3 where 
the Lemur catta population has been reduced as a result of hunting for food. Elsewhere such 
hunting as occurs appears to be mainly as a pastime rather than to provide a significant supply 
of proteim. It is thought likely that some animals, particularly lemurs, are caught alive for sale 

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Protected areas 

as pets. The reserve is seriously understaffed, with only two guards and a Chef de Reserve 
covering over 76 000 ha. 

SCIENTIFIC RESEARCH The area has been surveyed in the past (Paulian et al., 
1973); Propithecus has been the subject of a brief study by Richard at Hazafotsy (Parcel 2). 
There area is at present the subject of study by O'Connor. 

VISITOR AND SCIENTIFIC FACILITIES None. Access to Parcel 1 is difficult, although a 
road runs from Ranomafana to Isaka Ivondro on the eastern boundary. There are four paths, 
three of which follow the boundaries of the parcel with the fourth cutting through from east to 
west (Andonabe to Evasia). Parcel 2 is more accessible, from the road from Amboasary to 
Hazofotsy which continues to Ambatoabo, crossing the northern boundary of the parcel; this 
road is passable for most of the year and from it a path cuts south to Bevilany. Parcel 3 is 
easily accessible from R.N. 13 which runs from Fort Dauphin to Amboasary. There is a hut at 
Hazafotsy owned by the government agronomy service which has been used by visiting 
scientists. 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

O'Connor, S., Pidgeon, M., and Randria, Z. (1985). A conservation program for the 
Andohahela Reserve (Reserve Naturelle No. 11). Paper given at 'Seminaire Scientifique 
international sur l'etat de recherche sur l'equilibre des ecosystemes forestiers de 
Madagascar.' Antananarivo, October 1985. 

Paulian, R., Blanc, C, Guillaumet, J-L., Betsch, J-M., Griveaud, P. and Peyrieras, A. 
(1973). Etude des ecosystemes montagnards dans la region malagache. II Les chaines 
Anosyennes. Geomorphologie, climatologie et groupments vegeteaux. Campagne RCP 225 
1971-1972. Bull. Mus. Natn. Hist. Nat. Paris 3° ser. no. 118 Ecol. 1: 1-40. 

STAFF A Chef de Reserve (i.e. the Chef de Cantonnement Forestier et Poste R.N. 11), a 
Chef de Poste Est and a second guard, posted at Hazafotsy (see below). 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Protection of the reserve is coordinated 
by the Chef de Cantonnement Forestier et Poste R.N. 11, who is located at Amboasary-Sud. 
There are two guardposts, one at Eminiminy on the eastern side of Parcel 1 (Poste Est), the 
other at Hazafotsy on the northern boundary of Parcel 2. 

FAUNA 

Birds 

The following species have been recorded by O'Connor et al. (1985) in their surveys of all 

three parcels of the reserve; it is not regarded as a complete avifaunal list. 



Lophotibis cristata 
A viceda madagascariensis 
Polyboroides radiatus 
Accipiter madagascariensis 
Buteo brachypterus 
Falco newtoni (R) 
Margaroperdix madagarensis 
Turnix nigricollis 
Pterocles personatus 
Streptopelia picturata (R) 
Treron australis (R) 
Alectroenas madagascariensis 
Coracopsis nigra (R) 



Coracopsis vasa (R) 

Agapornis cana 

Coua gigas 

Coua reynaudii 

Coua cursor 

Coua cristata 

Coua caerulea 

Centropus toulou (R) 

Otus rutilus (R) 

Ninox superciliaris 

Caprimulgus madagascariensis (R) 

Alcedo vintsioides (R) 

Ipsidina madagascariensis 



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An environmental profile of Madagascar 



Eurystomus glaucurus (R) 
Leptosomus discolor (R) 
Motacilla flaviventris 
Coracina cinerea (R) 
Phyllastrephus sp 
Hypsi petes madagascariensis (R) 
Tylas eduardi 
Schetba rufa 
Vanga curvirostris 
Xenopirostris xenopirostris 
Falculea palliata 
Leptopterus viridis 
Hypositta corallirostris 



Copsychus albospecularis 
Neomixis viridis 
Cisticola cherina 
Newtonia brunneicauda 
Terpsiphone mutata (R) 
Nectarinia souimanga (R) 
Nectar inia notata (R) 
Zosterops maderspatana 
Lonchura nana (R) 
Ploceus nelicourvi 
Ploceus sakalava 
Foudia madagascariensis (R) 
Dicrurus forficatus (R) 



In addition there is a nineteenth century record of Brachypteracias squamiger, and Atelornis 
pittoides has been recorded from the Anosyennes hills. 

Mammals 

The following species have either been recorded by O'Connor and co-authors or identified 
by forest guards or villagers in the reserve. Numbers in parentheses indicate the parcel(s) in 
which the species has been recorded. A question mark indicates occurrence expected from 
known distribution of the species but not confirmed. 



Cheirogaleus medius [2,?3] 
Cheirogaleus major [ 1 ] 
Microcebus murinus [2,?3] 
Microcebus rufus [1] 
Phaner furcifer [?2] 
Avahi laniger [1] 
Le pi lemur mustelinus [1] 
Lepilemur leucopus [2,?3] 

Cryptoprocta ferox [1,2,?3] 
Galidia elegans [1] 
Salanoia concolor [1] 



Propithecus diadema [1] 
Propithecus verreauxi [1,2,?3] 
Daubentonia madagascariensis [1] 
Hapalemur griseus [ 1 ] 
Lemur fulvus [1] 
Lemur catta [1,2,3] 



Fossa fossa [1] 
Galidictis fasciata [?1] 



Amphibians 

No specific data, but the following amphibians are recorded from the Anosyennes chain of 
hills, the southern end of which comprises the R.N.I. d'Andohahela 



Anodontohyla boulengeri 
*Madecassophryne truebae 

Mantidactylus bertini 
*Mantidactylus grandisonae 

Mantidactylus redimitus 
*Boophis microtis 



* Anodontohyla rouxae 

*Microhyla palmata 
Mantidactylus elegans 
Mantidactylus microtympanum 
Mantidactylus tricinctus 



Information used in this account has been kindly provided by Sheila O'Connor and Mark 
Pidgeon. 



NAME Reserve Naturelle Integrate de MAROJEJY (No. 12). 
MANAGEMENT CATEGORY I (Strict Nature Reserve). 
LEGAL PROTECTION Total. 



■140- 



Protected areas 
DATE ESTABLISHED 3 January 1952. 

GEOGRAPHICAL LOCATION 14°18'-39'S, 49°33 , -52'E. To the north-west of Andapa in 
the province of Antseranana. 

ALTITUDE 90-2137 m. 

AREA 60 150 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The reserve includes the massif of Marojejy, and its principal 
foothills, notably Ambatosoratra, Ambodilahitra and Beondroka. The area comprises a very 
rugged massif chiefly composed of gneiss, which is divided into three main blocks. At around 
1100 m, the mid-slopes become increasingly steep, leading up to narrow quartzite ridges which 
precede the rocky escarpments encircling the west slopes of the massif. There is a wide range 
of microclimates. Rainfall on the eastern and south-eastern slopes is thought to reach or 
exceed 3000 mm a year, which would be the highest rainfall in Madagascar; measured rainfall 
at Andapa and Sambava is ca 2000 mm a year. Average temperatures at the lowest altitudes on 
the eastern side are approximately 22.3°C in July and 26.9°C in February. Winter temperatures 
on the summit of Marojejy (2133 m) are ca 1.5°C. 

VEGETATION Overall, plant species diversity is very high, with over 100 genera and 2000 
species recorded, several apparently endemic to the massif. Humbert (1955) has given a 
detailed description of the vegetation of the Marojejy Massif, divided into four altitudinal 
zones. The lowest zone, from 50 m to ca 800 m is high, dense, closed canopy rainforest, with a 
canopy height of 25-30 m and relatively clear horizontal stratification. Species diversity is 
very high. The most commonly represented families in the canopy are: Euphorbiaceae, 
Rubiaceae, Araliaceae, Ebenaceae (Diospyros), Sapindaceae, Sapotaceae, Anacardiaceae, 
Elaeocarpaceae (Echinocarpus), Lauraceae (Ocotea, Ravensara), Clusiaceae (Ochrocarpus), 
Myrtaceae, Burseraceae [Canarium), Moraceae, Bignoniaceae, Apocynaceae, Tiliaceae, 
Malpighiaceae, Monimiaceae, Flacourtiaceae, Loganiaceae. The intermediate stratum consists 
of small trees and large shrubs, mostly of the families Rubiaceae, Euphorbiaceae, Ochnaceae, 
Erthryoxylaceae, Myrsinaceae, Celastraceae, Violaceae, Flacourtiaceae. The ground layer is 
generally patchy, consisting of grasses and herbs, generally of the families Labiaceae, 
Acanthaceae, Gesneraceae, Melastomaceae, Balsaminaceae. Epiphytes are abundant. 

Transition from this stage to mid-altitude rainforest is made gradually, at around 800-900 m. 
This zone is characterised by a lower canopy height (18 to 25 m), with canopy trees generally 
branching lower down their trunks; the intermediate stratum tends to disappear, while the 
ground layer becomes denser and more varied. Although the species represented are generally 
different from those in the lower altitude forest, the families, and to some extent the genera, 
tend to be the same. Trees which are notably abundant or considered characteristic include 
species of Weinmannia, Apodocephala, Brachylaena and Podocarpus. Pteridophytes, particularly 
members of the Cyathaceae, are also abundant. 

Lichen or moss forest is best developed between 1450 and 1850 m, although it can be found as 
low as 1200 m altitude. Tree height is up to 6-10 m (max. 12 m), with trees often branching 
extensively from their bases. There is no intermediate stratum, but a dense and varied ground 
layer. Trees and shrubs mostly belong to the families Compositae (Vemonia, Senecio, 
Apodocephala, Psiadia), Lauraceae, Rubiaceae, Cunoniaceae {Weinmannia), Araliaceae 
(Cussonia), Euphorbiaceae (Uapaca, Acalypha, Croton), Rutaceae, Verbenaceae (Vitex, 
Clerodendron), Ericaceae (Agauria, Philippia), Sterculiaceae (Dombeya), Taxaceae (Podocarpus), 
Myricaceae (Myrica). Virtually monotypic stands of the bamboo Arundinaria marojejyensis are 
found on the least developed soils. Mosses and lichens are very abundant, both as ground 
cover and festooning trees and shrubs. Other epiphytes include ferns, species of Peperomia 
and orchids such as Bulbophyllum. 

Above 1850 m altitude is found vegetation which, depending on soil conditions and 
microclimate, has the appearance either of maquis or of heath. Shrubs chiefly belong to the 

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An environmental profile of Madagascar 

families Compositae, Ericaceae, Rubiaceae, Melastomaceae, Clusiaceae, Araliaceae, 
Euphorbiaceae, Myrtaceae; less numerous are members of the Cunoniaceae, Flacourtiaceae, 
Sapotaceae, Pittosporaceae, Sterculiaceae, Rutaceae, Verbenaceae, Vacciniaceae. Arundinaria 
marojejyensis forms scattered dense stands and tree ferns (Cyatheaceae) are found to ca 
2000 m altitude. Herbaceous plants consist largely of sedges (Cyperaceae) and grasses 
(Gramineae) and there are small marshes and swampy depressions with a distinctive flora. 

ZONING None. 

CONSERVATION MANAGEMENT The entire perimeter is marked by boundary stones 
connected by a largely overgrown footpath. There are four guards who each patrol a given 
sector of the boundary at about four-monthly intervals; however the guards lack equipment for 
prolonged foot patrols within the reserve. 

DISTURBANCES OR DEFICIENCIES The reserve is undermanned. Some tavy 

(slash-and-burn) cultivation is carried out in valleys which are rarely patrolled. The areas 
surrounding the reserve in the south and much of the west have been deforested to the 
boundary; the state of the northern and eastern boundaries is unknown. Before the reserve was 
created, coffee was grown at 400-500 m within the reserve boundaries; these areas are now 
slowly regenerating to native forest. The frequency of rain and storms provides effective 
protection of the reserve at higher altitudes. 

SCIENTIFIC RESEARCH One of the best studied massifs, first surveyed in 1933. 

VISITOR AND SCIENTIFIC FACILITIES None; there are no tourist paths or trails within 
the reserve, although a tractor track from Andapa to Doany provides access to the reserve in 
the west between 500 and 1000 m. 

PRINCIPAL REFERENCE MATERIAL 

Andriamampianina, J. and Peyrieras, A. (1972). Les reserves naturelles integrates de 
Madagascar. In: Comptes rendus de la Conference Internationale sur la Conservation de la 
Nature et de ses Ressources a Madagascar, Tananarive, Madagascar 7-11 octobre 1970. 
IUCN, Switzerland. 

Guillaumet, J-L., Betsch, J-M., Blanc, C, Morat, P., Peyrieras, A. and Paulian, R. (1975). 
Etude des ecosystemes montagnards dans la region malgache. III. Le Marojezy. IV. 
LTtremo et l'lbity. Geomorphologie, climatologie, faune et flore (Campagne RCP 225, 
1972-1973). Bull. Mus. Natn. Hist. Nat. (3). 309. (Ecol. generale): 25, 27-67. 

Humbert, H. (1955). Une merveille de la nature a Madagascar. Premiere exploration 
botanique du Massif de Marojejy et de ses satellites. Mem. Inst. sci. de Madagascar. 
Serie B. Tome VI. P.271. 

Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF - Protected areas 
programme in Madagascar. Unpd. report, 62 pp. 

STAFF An agent and 4 auxiliaries full-time. 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION Headquarters at Andapa, and guardposts 
at Doany and Ambalamanasy. 

FAUNA 



Birds 

Lophotibis cristata 
Aviceda madagascariensis 
Eutriorchis astur 
Polyboroides radiatus 
Accipiter henstii 
Accipiter madagascariensis 
Accipiter francesii (R) 



Buteo brachypterus 
Falco newtoni (R) 
Falco zoniventris 
Margaroperdix madagascarinus 
Turnix nig r icollis 
Dryolimnas cuvieri 
Mentocrex kioloides 



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Protected areas 



Sarothrura insularis 
Sarothrura watersi 
Streptopelia picturata (R) 
Treron austral is (R) 
Alectroenas madagascariensis 
Agapornis cana 
Coracopsis vasa (R) 
Coracopsis nigra (R) 
Coua caerulea 
Cuculus rochii 
Coua reynaudii 
Coua cristata 
Cent r opus toulou (R) 
Otus rutilus (R) 
Asio madagascariensis 
PNinox superciliaris 
Caprimulgus madagascariensis (R) 
Zoonavena grandidieri (R) 
Alcedo vintsioides (R) 
Ipsidina madagascariensis 
Eurystomus glaucurus (R) 
Leptosomus discolor (R) 
Br achy pier acias leptosomus 
Brachypteracias squamiger 
Atelornis crossleyi 
Philepitta castanea 
Neodrepanis coruscans 
Phedina borbonica (R) 
Motacilla flaviventris 
Coracina cinerea (R) 



Phyllastrephus madagascariensis 

Phyllasterphus zosterops 

Hypsi petes madagascariensis (R) 

Schetba rufa 

Vanga curvirostris 

Xenopirostris polleni 

Leptopterus chabert 

Leptopterus viridis 

Leptopterus madagascarinus (R) 

Oriolia bernieri 

Euryceros prevosti 

Hypositta corallirostris 

Tylas eduardi 

Monticola sharpei 

Nesillas typica 

Newtonia brunneicauda 

Neomixis viridis 

Neomixis tenella 

Terpsi phone mutata (R) 

Oxylabes madagascariensis 

Nectarinia souimanga (R) 

Nectarinia notala (R) 

Zosterops maderaspatana 

Lonchura nana (R) 

Ploceus nelicourvi 

Foudia madagascariensis (R) 

Foudia omissa 

Saroglossa aurata 

Dicrurus forficatus (R) 



Mammals 

Hapalemur griseus 

Propithecus diadema 

Lemur fulvus 

Lemur macaco 

Varecia variegata reported 

Cryptoprocta ferox 



Galidia elegans 
Hemicentetes semispinosus 
Microgale talazaci 
Tenrec ecaudatus 
Microcebus rufus 
Pteropus rufus 



Amphibians 

*Mantipus minutus 
Plethodontohyla notosticta 
Rhombophryne testudo 
Stumpffia psologlossa 

*Stumpffia tridactyla 
Laurentomantis horrida 
Mantidactylus bicalcaratus 
Mantidactylus lugubris 
Mantidactylus redimitus 



*Mantipus serratopal pebrosus 

Plethodontohyla ocellata 
*Stumpffia grandis 
*Stumpffia roseifemoralis 
Dyscophus insularis 
Mantidactylus asper 
*Mantidactylus klemmeri 
*Mantidactylus pseudoasper 



Reptiles 

Zonosaurus aff. rufipes 
*Brookesia griveaudi 
Chamaeleo gastrotaenia 
Chamaeloe nasutus 
Brookesia betschi 
Chamaeleo aff. malthe 
Pararhadinea melanogaster 



*Brookesia karchei 
Chamaeleo peyrierasi 
Chamaeleo globifer 
Chamaeleo bifidus 
Chamaeleo aff. brevicornis 
Brookesia aff. minima 
Liopholidophis stumpffi 



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An environmental profile of Madagascar 



Nonmarine Molluscs 

Those marked + have definitely been recorded within the reserve. 

Acroptychia metablata + Ampelita gaudens + 

Tropidophora tricar inat a * Ampelita globulus + 

Tropidophora zonala Ampelita lamarei + 

Helicophanta amphibulima + Ampelita perampla + 

Kalidos oleatus + *Malagrion paenelimax 
Macrochlamys stumpfii + 



NAME Reserve Special Botanique d'AMBOHITANTELY. 

MANAGEMENT CATEGORY IV (Managed Nature Reserve). 

LEGAL PROTECTION Protection of all resources within the reserve is total. Access is not 
restricted by the establishing decree. 

DATE ESTABLISHED 12 February 1982, upgraded from Foret classee to a Reserve Speciale 
Botanique. 

GEOGRAPHICAL LOCATION Ca 18°08'-18°13'S, 47 18'-47°21'E; on the Tampoketsa 
d'Ankazobe some 80 km north of Antananarivo. 

ALTITUDE 1200-1650 m. 

AREA 5600 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The reserve consists of 'tampoketsa', that is levelled off remains of 
ancient erosion surfaces at high altitude, generally forming highly dissected plateaux bordered 
by steep escarpments. The formation is assumed to be late Cretaceous in origin. Altitude of 
the tampoketsa is 1600 m, though the forest descends to around 1450 m. Rainfall averages 
around 1500 mm per annum, with a marked wet season from November to March. Mean 
maximum temperature is 26°C, minimum 12°C. 

VEGETATION The area contains one of the few remaining vestiges of the central plateau 
forest. In 1964, the forest consisted of a single tract of ca 2000 ha on the eastern slope of the 
tampoketsa, with an additional 1000 ha of small scattered fragments mainly at the heads of 
valleys on the tampoketsa itself. The forest has very close floristic affinities with the eastern 
rain forest, especially at lower altitudes, where it appears to be more or less primary. 

ZONING None. 

CONSERVATION MANAGEMENT Ambohitantely is the focus for IUCN/WWF Project 1912, 
Protection and Management of Ambohitantely Forest Reserve. The reserve was first set up on 
the initiative of the Direction des Eaux et Forets with the support of WWF. This project, 
under the responsibility of the Plant Biology and Biochemical Service of Antananarivo 
University, is mainly scientifically orientated in support of management. Work is in progress 
on an inventory of the flora. The Plant Biology and Biochemical Service made two visits to the 
area in 1982 and three in 1983, with the aid of the WWF Representation, which provided the 
vehicle and fuel. WWF will also support the development of this newly established reserve. 
The area could be particularly valuable as a training area because of its proximity to 
Antananarivo. Studies made here will help in developing plans to reafforest the tampoketsa 
region using native trees, and will form the basis of a public conservation education campaign. 



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Protected areas 

DISTURBANCES OR DEFICIENCIES The most serious danger is fire, but boundaries also 
need to be marked, guards installed, access controlled, and paths maintained. 

SCIENTIFIC RESEARCH Studies of the fauna and flora, carried out by Antananarivo 
University, under IUCN/WWF Project 1912, concentrate on plants of medicinal, ornamental, or 
possible economic use, and on endangered or rare species. Studies also concentrate on the 
effect of fire on the vegetation. 

VISITOR AND SCIENTIFIC FACILITIES There are no facilities within the reserve, but there 
are nearby facilities at Antananarivo. 

PRINCIPAL REFERENCE MATERIAL 

IUCN/WWF Project 1912. Protection and Management of Ambohitantely Forest Reserve. 
Bastian, G. (1964). La foret d' Ambohitantely, Madagascar. Revue de Geographie 
5: 1-42. 

STAFF No information. 

BUDGET The reserve has received funding from the WWF direction a Madagascar. 

LOCAL PARK OR RESERVE ADMINISTRATION There is a forest station at Manankazo. 

FAUNA 

Mammals 

Lemur fulvus 

Reptiles 

The following records were provided by C. Raxworthy (in litt., 4.12.86). 

Chamaeleo nasutus Phesluma lineata 

Chamaeleo parsonii 

Amphibians 

Mantidactylus peraccae Platypelis pollicaris 

Mantidactylus punctatus Plethodontohyla laevis 

Nonmarine Molluscs 

Vitrina madagascariensis 

Nonmarine Crustacea 
Decapoda 

Hydrothelphusa humbloti 



NAME Reserve Speciale de BEZA MAHAFALY 

MANAGEMENT CATEGORY IV (Managed Nature Reserve). 

LEGAL PROTECTION Total. 

DATE ESTABLISHED 1979. Inaugurated November 1985. 

GEOGRAPHICAL LOCATION Just west of the Sakamena River, about 35 km north-east of 
Betioky-Sud; ca 23°30'S, 44°40'E. The reserve is divided into two non-contiguous parcels, one 
lying along the Sakamena River, the second some 5 km west of the Sakamena. 

ALTITUDE Ca 100-200 m. 

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An environmental profile of Madagascar 

AREA 600 ha in two parcels, one of 100 ha, the second of 500 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The first (100 ha) parcel borders the Sakamena River, which 
normally contains water during the rainy season, from November or December to March; for 
the rest of the year it is a dry sandy river bed (Richard el al., 1985). 

VEGETATION The first parcel consists of low gallery forest, dominated by Tamarindus 
indica; the second consists of spiny forest dominated by Alluaudia procera with other members 
of the Didiereacae and Euphorbiaceae (Richard et al., 1985). 

ZONING None apart from the two defined areas. 

CONSERVATION MANAGEMENT The smaller parcel is bounded by a barbed-wire fence, 
erected in 1979; the larger has a 3 m swathe cut around it to delineate the boundaries. Opuntia 
has been planted to provide an effective barrier. Grids of trails have been cut within the 
reserve, these being 100 m-to-a-side in the smaller parcel and 500 m-to-a-side in the larger 
(Richard et al., 1985). 

DISTURBANCES OR DEFICIENCIES The reserve appears to be well protected at present. 
Cattle and goats formerly ranged throughout the forests at Beza Mahafaly; since 1979 these 
have been excluded from the smaller parcel by the boundary fence, though it is not clear if 
they still enter the larger (Richard et al., 1985). 

SCIENTIFIC RESEARCH The Beza Mahafaly project is based on an Inter-University Accord 
between the University of Madagascar, Yale University and Washington University. One of its 
principal roles is to provide a site for research on the flora and fauna of the south-west of 
Madagascar, and on the relations between the Madagascan people and the natural environment. 
The importance of baseline survey work as a preliminary to more detailed studies has been 
emphasised (Richard et al., 1985). Up to the present, more or less detailed inventories of 
primates, insectivores and rodents, birds and insects, particularly Hymenoptera, have been 
carried out (see Fauna below) along with a study of the structure and composition of the 
vegetation inside and outside the reserve. This last study is intended to give information on 
the regeneration of the natural vegetation and the impact of livestock grazing on this, with the 
reserve, being fenced, effectively acting as a large scale exclosure. A more detailed study of 
the demography and behaviour of Propithecus verreauxi in the larger parcel of the reserve was 
begun in 1984. Plans for further research include: a study of the ethnomedicine of the region, 
to be expanded to a general ethnobotanical survey of the area; extending the study of forest 
structure, diversity and regeneration to the second parcel; establishment of an on-site 
herbarium; exploration of the phenology and pollination ecology of dominant tree and shrub 
species in the two parcels; a study of the behavioural ecology of Lemur catta; research on the 
reptile community of the reserve with a detailed study of Geochelone radiata (Richard et al., 
1985). 

VISITOR AND SCIENTIFIC FACILITIES Huts for equipment and cooking. 

PRINCIPAL REFERENCE MATERIAL 

Rakotomanga, P., Richard, A.F. and Sussman, R.W. (1985). Beza Mahafaly. Formation et 
Mesures pour la conservation. Paper given at 'Seminaire Scientifique international sur 
l'etat de recherche sur l'equilibre des ecosystemes forestiers de Madagascar.' 
Antananarivo, October 1985. 

Richard, A.F., Rakotomanga, P. and Sussman, R.W. (1985). Beza Mahafaly: recherches 
fondamentales et appliquees. Paper given at 'Seminaire Scientifique international sur l'etat 
de recherche sur l'equilibre des ecosystemes forestiers de Madagascar.' Antananarivo, 
October 1985. 

STAFF One chief warden, five permanent guards plus auxiliary guards. 



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Protected areas 

BUDGET The Beza Mahafaly project has received financial support from WWF-US since 
1980. 

LOCAL PARK OR RESERVE ADMINISTRATION Reserve headquarters are at Betioky-Sud. 

FAUNA 

Birds 

The following species have been recorded by Randrianasolo and Pidgeon {fide Richard et 
al., 1985) and updated by Pidgeon (pers. comm., 1986). 



Polyboroides radiatus 
Accipiter madagascariensis 
Accipiter francesii (R) 
Buteo brachypterus 
Falco newtoni (R) 
Margaroperdix madagarensis 
Turnix nigricollis 
Dryolimnas cuvieri 
Streptopelia picturata (R) 
Coracopsis nigra (R) 
Coracopsis vasa (R) 
Agapornis cana 
Coua gigas 
Coua cursor 
Coua cristata 
Centropus toulou (R) 
Otus rutilus (R) 
Ninox superciliaris 
Alcedo vintsioides (R) 

Mammals 

Propithecus verreauxi 
Lepilemur leucopus 
Lemur catta 
Microcebus murinus 
Cryptoprocta ferox 



Leptosomus discolor (R) 
Coracina cinerea (R) 
Hypsi petes madagascariensis (R) 
Vanga curvirostris 
Xenopirostris xenopirostris 
Falculea palliata 
Leptopterus chabert 
Leptopterus viridis 
Eurystomus glaucurus (R) 
Copsychus albospecularis 
Neomixis tenella 
Neomixis striatigula 
Newtonia brunneicauda 
Terpsiphone mutata (R) 
Nectarinia soui manga (R) 
Zosterops maderaspatana 
Ploceus sakalava 
Foudia madagascariensis (R) 
Dicrurus forficatus (R) 



Echinops telfairi 
Setifer setosus 
Geogale aurita 
Tenrec ecaudatus 
Suncus madagascariensis 



Cheirogaleus medius has been recorded within 1 km of the reserve. This is the only known 
site where Geogale aurita is common (M. Nicoll, in litt., 28.10.86). 



Reptiles 

The following species have been recorded by C. Raxworthy (in. 
Pidgeon and S. O'Connor (pers. comm., 10.10.86). 



UtL, 4.12.86) and M. 



Geochelone radiata 
Hemidactylus mabouia 
Homopholis sakalava 
Phelsuma mutabilis 
Geckolepis typica 
Paroedura bastardi 
Paroedura pictus 

Tracheloptychus madagascariensis 
Mabuya gravenhorsti 



Mabuya elegans 
Mabuya aureopunctata 
Chamaeleo verrucosus 
Leioheterodon geayi 
Leioheterodon madagascariensis 
Ithycyphus mini at us 
Acrantophis dumerilii 
Chalaradon madagascariensis 
Erymnochelys madagascariensis 



Much of the information in this account has been kindly provided by Sheila O'Connor and 
Mark Pidgeon. 



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An environmental profile of Madagascar 

NAME Reserve Speciale de NOSY MANGABE 

MANAGEMENT CATEGORY IV (Managed Nature Reserve). 

LEGAL PROTECTION Protection of all resources within the reserve is total. Access is not 
restricted by the establishing decree. 

DATE ESTABLISHED 14 December 1965 by Decree No. 65-795. 

GEOGRAPHICAL LOCATION 15°25'S, 49°45'E. A small island situated to the east of 
Maroantsetra, 6 km off the coast of Madagascar in the Bay of Antongil. 

ALTITUDE From sea level to 331 m. 

AREA 520 ha (the whole island). 

LAND TENURE Government owned. 

PHYSICAL FEATURES Cretaceous limestone island, with very rugged topography. 

VEGETATION The island has a typical east coast rainforest vegetation including species 
of Canarium, Ocotea and Ravensara, along with many palms and ferns. Much of the forest is 
secondary. 

ZONING None. 

CONSERVATION MANAGEMENT Aye-aye Daubentonia Madagascariensis were introduced 
onto the island in 1966. Other than this, relatively little active management appears to have 
been carried out, although the island does not appear to be unduly disturbed. A boat with 
outboard motor is available for patrols and there is a building which could serve as a laboratory 
on the island. 

DISTURBANCES OR DEFICIENCIES The island can only support limited tourism and this 
must be more strictly controlled. It has been suggested (IUCN Project 1953) that no more 
buildings be constructed on the island. Planned staff housing should be built on the adjacent 
mainland. Any manipulation of the habitat should be strictly minimised. There is a 
fishermen's hut at one end of the island which is in constant use. 

SCIENTIFIC RESEARCH A study of Varecia variegata was due to begin in autumn 1986. 

VISITOR AND SCIENTIFIC FACILITIES A one room laboratory. 

PRINCIPAL REFERENCE MATERIAL 

IUCN/WWF Project 1953. 

STAFF Two agents and two auxiliaries. 

BUDGET Salaries paid by the government. WWF has provided funds for management. 

LOCAL PARK OR RESERVE ADMINISTRATION No information. 

FAUNA 

Birds 

Records are from O. Langrand (unpublished data, in litt., 28.10.86). 

Accipiter francesii (R) Coracopsis nigra (R) 

Buteo brachypterus Cuculus rochii (R) 

Dryolimnas cuvieri Centropus toulou (R) 

Streplopelia picturata ( R) Alcedo vintsioides (R) 

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Protected areas 

Ipsidina madagascariensis Newtonia amphichroa 

Leptosomus discolor (R) Terpsi phone mutata (R) 

Motacilla flaviventris Nectarinia souimanga (R) 

Hypsi petes madagascariensis (R) Nectarinia notata (R) 

Leptopterus chabert Zosterops maderaspatana 

Copsychus albospecularis Ploceus nelicourvi 

Nesillas typica Dicrurus forficatus (R) 

Mammals 

Daubentonia madagascariensis Varecia variegata 

Microcebus rufus Setifer setosus 

Lemur fulvus Oryzorictes sp. 

Pteropus rufus is present on islands 1 km distant (M. Nicoll, in litt., 28.10.86). 

Amphibians 

?* Mantella laevigata ?* Mantidactylus webbi 

* Boophis leucomaculatus Dyscophus antongili 

Reptiles 

The following species, excepting Liopholidophis thieli, have been recorded by Q. Bloxam (in 
litt., 23.07.86) and C. Raxworthy (in litt., 4.12.86). The record for Pseudoxyrhopus heterurus 
is the first definite locality record for this species. 

Sanzinia madagascariensis Uroplatus fimbriatus 

Liopholidophis thieli Chamaeleo oustaleti 

Pseudoxyrhopus heterurus Chamaeleo pardalis 

Homopholis antongilensis Brookesia peyrierasi 

Phelsuma guttata Zonosaurus aeneus 

Paroedura androyensis Zonosaurus madagascariensis 

Ebenavia inunguis Amphiglossus sp. 



NAME Reserve de faune de PERINET-ANALAMAZOATRA. 

MANAGEMENT CATEGORY IV (Managed Nature Reserve). 

LEGAL PROTECTION All resources within the reserve are totally protected. Access is 
theoretically not controlled by the decree setting up the reserve, although a permit, issued by 
the Direction des Eaux et Forets, is required for entry. 

DATE ESTABLISHED 21 June 1970. 

GEOGRAPHICAL LOCATION 100 km east of Antananarivo to the east of Moramanga; 
18°28'S, 48°28'E. 

ALTITUDE 930-1040 m. 

AREA 810 ha. 

LAND TENURE Government land. 

PHYSICAL FEATURES The reserve lies within a crystalline massif with rugged topography; 
soils are principally lateritic. Annual rainfall is ca 1700 mm, with most in January and least in 
October. Mean monthly temperature varies from 14°C in August to 24°C in January; cyclones 
may occur between November and March. 



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An environmental profile of Madagascar 

VEGETATION Medium altitude tropical moist forest; characteristic canopy genera 

include Weinmannia, Tambourissa, Symphonia, Dalbergia, Ravensara and Vernonia. The 
understorey strata are particularly dense and include representatives of Cyathea, Dypsis, 
Plantago, Smilax, Rubus, Alchemilla and Sanicula. Epiphytes, including orchids 
(especially Bulbophyllum) and Rhipsalis, are abundant. 

ZONING None. 

CONSERVATION MANAGEMENT The reserve is well marked on the ground and there are 
numerous well-used trails. The reserve is generally left undisturbed apart from path clearing 
activities; patrols are carried out on an ad hoc basis and only occasionally penetrate the furthest 
reaches of the reserve. A private organization, the Friends of the Reserves of Andasibe 
(Perinet), has recently been formed and could play an active role in assisting in the 
maintenance of the reserve. 

DISTURBANCES OR DEFICIENCIES The reserve is too small to protect this forest type 
adequately. Tavy (slash-and-burn) is the principal cultivation technique used in the region; 
with the increasing human population it is becoming unsustainable and poses a severe 
long-term threat. The reserve is being encroached from the south and east by tavy, though is 
still buffered to the north and west by native and plantation forests. There is evidence of 
hunting and exploitation of hardwoods within the reserve. There is also substantial collection 
of animals to supply both the national and overseas pet trades; animals involved 
include Phelsuma, Chamaeleo and Mantidactylus spp, Sanzinia madagascariensis, Microcebus 
rufus, Hapalemur griseus and Lemur fulvus. 

SCIENTIFIC RESEARCH Many studies have been carried out on the fauna and flora of the 
reserve. 

VISITOR AND SCIENTIFIC FACILITIES The reserve is easily accessible from Antananarivo 
and Toamasina, either by rail or by road; the latter passes alongside the reserve. The forest 
station associated with the reserve has accomodation, though this is in a poor state of repair. 

PRINCIPAL REFERENCE MATERIAL 

Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF - Protected areas 
programme in Madagascar. Unpd. report, 62 pp. 

STAFF The Chef de Station Forestiere is responsible for the reserve; there are also two 
labourers. 

BUDGET Salaries paid by the government. 

LOCAL PARK OR RESERVE ADMINISTRATION The reserve is under the responsibility of 
the Service de la Protection de la Nature, Direction des Eaux et Forets, B.P. 243, 
Antananarivo. Local responsiblity lies with the Chef de Station Forestiere d'Analamazoatra. 

FAUNA 



Birds 

Records are from Dee (in press) (see part V.I.), amended by M. Pidgeon, O. 
Thompson, J. Thorsen, J. Ganzhorn and T. Moermond (unpublished data). 



Langrand, P. 



Tachybaptus pelzelnii 
Ardeola idae (R) 
Lophotibis cristata 
Anas melleri 

A viced a madagascariensis 
Eutriorchis astur 
Polyboroides radiatus 
Accipiter madagascariensis 
Accipiter henstii 



Accipiter francesii (R) 
Buteo brachypterus 
Falco newtoni (R) 
Falco zoniventris 
Margaroperdix madagarensis 
Mesitornis unicolor (probable) 
Dryolimnas cuvieri 
Canirallus kioloides 
Sarothrura insularis 



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Protected areas 



Sarothrura watersi 

Streptopelia picturata (R) 

Treron austral is (R) 

Alectroenas madagascariensis 

Coracopsis vasa (R) 

Coracopsis nigra (R) 

Cuculus rochii 

Coua cristata 

Coua caerulea 

Coua serriana 

Coua reynaudii 

Centropus toulou (R) 

Tyto soumagnei 

Otus rutilus (R) 

Asio madagascariensis 

Caprimulgus enarratus 

Caprimulgus madagascariensis (R) 

Zoonavena grandidieri (R) 

Alee do vintsioides (R) 

Ipsidina madagascariensis 

Eurystomus glaucurus (R) 

Brachypteracias leptosomus 

Atelornis pittoides 

Atelornis crossleyi 

Leptosomus discolor (R) 

Philepilta castanea 

Neodrepanis coruscans 

Neodrepanis hypoxantha 

Motacilla flaviventris 

Coracina cinerea (R) 

Phyllastrephus madagascariensis 

Phyllastrephus zosterops 

Phyllastrephus tenebrosus 

Hypsi petes madagascariensis (R) 

Tylas eduardi 

Calicalicus madagascariensis 



Vanga curvirostris 
Xenopirostris polleni 
Leptopterus chabert 
Leptopterus madagascarinus (R) 
Leptopterus viridis 
Hypositta corallirostris 
Copsychus albospecularis 
Monticola sharpei 
Neomixis viridis 
Neomixis striatigula 
Neomixis tenella 
Hartertula flavoriridis 
Oxylabes madagascariensis 
Mystacornis crossleyi 
Acrocephalus newtoni 
Crossleyia xanthophrys 
Nesillas typica 
Cisticola cherina 
Dromaeocercus brunneus 
Dromaeocercus seebohmi 
Randia pseudozosterops 
Newtonia amphichroa 
Newtonia brunneicauda 
Newtonia fanovanae 
Pseudobias wardi 
Terpsi phone mutata (R) 
Nectarinia soui manga (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Lonchura nana (R) 
Ploceus nelicourvi 
Foudia madagascariensis (R) 
Foudia omissa 
Saroglossa aurata 
Dicrurus forficatus (R) 



Neodrepanis hypoxantha has been recorded just outside the reserve. 



Mammals 

Cheirogaleus major 

Microcebus rufus 

Avahi laniger 

Indri indri 

Daubentonia madagascariensis 

Hapalemur griseus 

Lemur fulvus 

Lemur rubriventer 

Lepilemur microdon 

Hemicentetes semispinosus 

Microgale taiva 

Microgale thomasi 

Microgale talazaci 



Microgale melanorrhachis 
Microgale pusilla 
Microgale gracilis 
Oryzorictes hova 
Setifer setosus 
Tenrec ecaudatus 
Suncus madagascariensis 
Brachytarsomys albicauda 
Eliurus myoxinus 
Eliurus minor 
Gymnuromys roberti 
Nesomys rufus 



Varecia variegata has been recorded in the vicinity, although there are no records from the 
reserve itself; Propithecus diadema previously occurred in the reserve but no longer does so, 
although it is still present in the area. 



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An environmental profile of Madagascar 



Amphibians 

Anodontohyla boulengeri 
Platyhyla grandis 
Platypelis tubifera 
?*Mantidactylus acuticeps 

*Mantidactylus eiselti 
Mantidactylus liber 
Mantidactylus pulcher 
Boophis difficilis 
Boophis granulosus 
Boophis idae 
Boophis paulianus 

*Boophis reticulatus 

*Boophis viridis 



*Paracophyla tuberculata 
Platypelis pollicaris 

*Mantella aurantica 
Mantidactylus blommersae 
Mantidactylus flavobrunneus 
Mantidactylus o pi pari s 
Mantidactylus tornieri 
Boophis erythrodactylus 
Boophis hilleni 
Boophis miniatus 
Boophis rappoides 
Boophis untersteini 



Reptiles 

Several of the following are records provided by C. Raxworthy (in litt., 4.12.86). 



*Lygodactylus guibei 
*Phelsuma flavigularis 
Phelsuma lineata 
Uroplatus fimbriatus 
Brookesia theili 
*Brookesia therezieni 
Chamaeleo nasutus 
Chamaeleo parsonii 

Nonmarine Crustacea 
Isopoda: 

Suarezia differens 
Philoscia reducta 
Bethalus bipunctatus 
Calmanesia erinaceus 



Chamaeleo willsii 
Amphiglossus melanopleura 
Pararhadinea albignaci 
Pararhadinea melanogaster 
Micropisthodon ochracheus 
Liopholidophis stumpffi 
Liopholidophis thieli 



Didima humilis 
Armadillo otion 
Akermania hystrix 



NAME Reserve d'ANALABE 

MANAGEMENT CATEGORY IV (Managed Nature Reserve). 

LEGAL PROTECTION Private land. Status unknown. 

DATE ESTABLISHED Reserve management and status under revision since 1984. 

GEOGRAPHICAL LOCATION The centre of the reserve is at 19°29'S, 44°34'E; 60 km north 
of Morondava. The reserve is bounded to the north by sisal plantations around the village of 
Beroboka sud; there is no other clearly defined limit. The eastern edge of the reserve is close 
to a permanent lake and marsh system. 

ALTITUDE Entirely below 100 m above sea level. 

AREA Unclear; somewhere in the range 2000 to 12 000 ha. 

LAND TENURE Private land owned by the de Heaulme family. 

PHYSICAL FEATURES The reserve lies on the Morondava coastal plain; the land is flat with 
depressions forming seasonal or permanent marshes or lakes. Soil is sandy and water retention 
following rains is low. Annual rainfall is in the region of 700-1500 m, almost all of which falls 
from November to February, with most in January. Mean annual maximum temperature is 






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Protected areas 

31.1°C, mean annual minimum temperature 18.9°C. The reserve extends to the Mozambique 
Channel and includes some mangrove and beach features. There is evidence of many seasonal 
stream beds. 

VEGETATION Mainly dry western deciduous forest. Notable trees include 

baobab Adansonia grandidieri and tamarind Tamarindus indica. There is also thorny and 
succulent bush and scrub. There are many lianes and the undergrowth can be very dense. 
Average tree height is low, with the canopy generally at 10-13 m, though baobabs can reach 
15 m. 

ZONING The reserve is split by the remains of a sisal plantation. Plans have been drawn up 
by the owner, J de Heaulme, to divide the reserve into three parcels. One would be a strict 
nature reserve, the second a parcel for scientific research and the third for tourism. 

CONSERVATION MANAGEMENT A reserve manager was present until early 1986; currently 
(1987) there is no effective management. 

DISTURBANCES OR DEFICIENCIES The reserve is unprotected at present. Incursions for 
tavy cultivation are being made and there is a high risk of damage from bush fires which burn 
surrounding forest annually. Tracks for oil exploration have been cut through the forest on a 
5 km grid system; these have made the reserve very vulnerable to exploitation, particularly for 
fuel wood. Fuel wood collection does not appear to be very extensive at present but is 
expected to increase. Hunting, principally for Tenrec ecaudatus and Hypogeomys antimena, 
occurs throughout the reserve; its current impact is unknown, though may not be very great (D. 
Curl in litt. to S. O'Connor, 27.7.86). There is evidence of burning along the roads and burned 
patches for small agricultural plots were seen 100 m off the road; wood is taken out of the 
reserve. 

SCIENTIFIC RESEARCH Little at present. 

VISITOR AND SCIENTIFIC FACILITIES Under consideration. Plans exist to renovate 
existing houses and plantation buildings to provide lodges and a conservation education centre; 
a research centre may also be included in the development plans. 

PRINCIPAL REFERENCE MATERIAL 

Nicoll, M.E. and Langrand, O. (1987). Report on the first phase of WWF - Protected areas 
programme in Madagascar. Unpd. report, 62 pp. 

STAFF None at present. 

BUDGET Unknown. 

LOCAL PARK OR RESERVE ADMINISTRATION None; the reserve is owned by J. de 
Heaulme, B.P. 37, Taolanaro. 

FAUNA 

Birds 

Records are from O. Langrand (unpublished data, in litt., 28.10.86) and S. O'Connor and M. 
Pidgeon (pers. comm., 10.10.86). 

Lophotibis cristata Margaroperdix madagarensis 

Anas bernieri Mesitornis unicolor 

Aviceda madagascariensis Turnix nigricollis 

Haliaeetus vociferoides Dryolimnas cuvieri 

Polyboroides radiatus Charadrius thoracicus 

Accipiter francesii (R) Pterocles personatus 

Buteo brachypterus Streptopelia picturata (R) 

Falco newtoni (R) Treron australis 

Falco zoniventris Coracopsis vasa (R) 

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An environmental profile of Madagascar 



Coracopsis nigra (R) 
Agapornis cana 
Cuculus rochii 
Coua gigas 
Coua coquereli 
Coua cristata 
Coua ruficeps 
Centropus toulou (R) 
Otus rutilus (R) 
Ninox superciliaris 
Asio madagascariensis 
Caprimulgus madagascariensis (R) 
Zoonavena grandidieri (R) 
Alcedo vintsioides (R) 
Leptopterus madagascarinus (R) 
Eurystomus glaucurus (R) 
Leptosomus discolor (R) 
Mirafra hova 
Phedina borbonica (R) 
Motacilla flaviventris 
Coracina cinerea (R) 



Phyllastrephus madagascariensis 
Hypsi petes madagascariensis (R) 
Vanga curvirostris 
Falculea palliata 
Leptopterus viridis 
Leptopterus chabert 
Leptopterus madagascarinus (R) 
Copsychus albospecularis 
Acrocephalus newtoni 
Nesillas typica 
Cisticola cherina 
Newtonia brunneicauda 
Neomixis tenella 
Terpsiphone mutata (R) 
Nectar inia notata (R) 
Nectar inia soui manga (R) 
Zosterops maderaspatana 
Ploceus nelicourvi 
Ploceus sakalava 
Lonchura nana (R) 
Dicrurus forficatus (R) 



Mammals 

Lemur fulvus 
Lepilemur ruficaudatus 
Microcebus murinus 
Microcebus coquereli 
Mungotictis striata 
Tenrec ecaudatus 



Propithecus verreauxi 
Phaner furcifer 
Cheirogaleus medius 
Hypogeomys antimena 
Cryptoprocta ferox 



Reptiles 

Pyxis planicauda 
Erymnochelys madagascariensis 
Oplurus cuvieri 
Acrantophis dumerilii 



Leioheterodon madagascariensis 
?Leioheterodon geayi 
Chamaeleo verrucosus 
Chalaradon madagascariensis 



NAME BERENTY Reserve 

MANAGEMENT CATEGORY IV (Managed Nature Reserve). 

LEGAL PROTECTION The reserve is privately owned, though is held in a trust which 
should ensure long-term protection. 

DATE ESTABLISHED 1930s; precise date unknown. 

GEOGRAPHICAL LOCATION 24°50'S, 46°20'E. Near Amboasary and 80 km from 
Taolanaro (Fort Dauphin). 

ALTITUDE Ca 0-30 m. 

AREA Berenty Reserve is composed of 5 parcels, comprising some 250-265 ha in total. 
Parcel 1 (known as Malaza) is 200 ha, parcel 2 ca 20 ha, parcel 3 ca 12 ha, parcel 4 ca 2 ha, 
parcel 5 (Anjapolo) is 20-30 ha. In addition the 97 ha forest of Bealoka some 7 km north of 
Berenty is to be incorporated into the reserve system. 

LAND TENURE Owned by the de Heaulme family. 



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Protected areas 

PHYSICAL FEATURES The reserve is situated in a sisal plantation and is bordered by the 
Mandrare River. 

VEGETATION Parcel 1 consists of a corridor of spiny forest dominated by Euphorbiaceae, 
Didiereacae and crassulids, with gallery forest with Acacia and Tamarindus close to the river; 
parcels 2 and 5 consist of spiny forest; parcel 3 is a planted forest of Pithecelobium dulce mixed 
with old Acacia and Tamarind trees; parcel 4 is a 2 ha sacred spiny forest with tombs. Bealoka 
consists of degraded gallery forest. 

ZONING None. 

CONSERVATION MANAGEMENT Reportedly well protected, being fenced and guarded. 
Little active management takes place at present. 

DISTURBANCES AND DEFICIENCIES Principal problems are: river flooding and river bank 
erosion, with areas of river bank up to 20 m wide having been lost from parts of the reserve 
since the 1960s; mature tree die-off, mainly affecting Tamarindus, Acacia and Nestina, the 
causes of which are unknown; poor regeneration in many parts of the forest, due to invasion of 
the rubber vine Cissus quadrangular is. 



SCIENTIFIC RESEARCH 

most recently by O'Connor. 



Work has been carried out on the lemurs since the early 1960s, 



VISITOR AND SCIENTIFIC FACILITIES 

entrance fee is charged. 



There are guest houses and a museum. A small 



PRINCIPAL REFERENCE MATERIAL 

Jolly, A. (1966). Lemur behavior. Chicago University Press. 

Jolly, A., Oliver, W.L.R. and O'Connor, S.M. (1982). Population and troop ranges of 

Lemur catta and Lemur fulvus at Berenty, Madagascar: 1980 census. Folia primatologica 

39(1-2): 115-123. 
Mertl-Millhollen, A.S., Gustafson, H.L., Budnitz, N., Dainis, K. and Jolly, A. (1979). 

Population and territory stability of the Lemur catta at Berenty, Madagascar. Folia 

primatologica 31: 106-22. 

STAFF The whole reserve is under the control of the manager of the sisal concession in 
which it is situated. There are four guards in parcel 1, two in parcel 2, one each in parcel 3 
and 5, and two at Bealoka. 

BUDGET Not known. 

LOCAL PARK OR RESERVE ADMINISTRATION See 'staff above. 

FAUNA 



Birds 

Most of the following species were recorded at Berenty during the period September 1983 - 
May 1986 by M.S. Pidgeon; additional records were provided by O. Langrand (unpublished 
data, in litt., 28.10.86). 



Ardea humbloti 
?Aviceda madagascariensis 
Polyboroides radiatus 
Accipiter madagascariensis 
Accipiter francesii (R) 
Buteo brachypterus 
Falco newtoni (R) 
Falco zoniventris 
Turnix nigricollis 
Dryolimnas cuvieri 



Pterocles personatus 
Streptopelia picturata (R) 
Treron australis 
Coracopsis nigra (R) 
Coracopsis vasa (R) 
Agapornis cana 
Cuculus rochii 
Coua gigas 
Coua cristata 
Cent ro pus toulou (R) 



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An environmental profile of Madagascar 



Otus rutilus (R) 
Ninox superciliaris 
Caprimulgus madagascariensis 
Zoonavena grandidieri (R) 
Alcedo vintsioides (R) 
Ipsidina madagascariensis 
Leptosomus discolor (R) 
Eurystomus glaucurus (R) 
Mirafra hova 
Phedina borbonica (R) 
Motacilla flaviventris 
Coracina cinerea (R) 
Hypsi petes madagascariensis (R) 
Calicalicus madagascariensis 
Xenopirostris xenopirostris 
Vanga curvirostris 
Falculea palliata 
Leptopterus chabert 



Leptopterus viridis 

Leptopterus madagascarinus (R) 

Copsychus albospecularis 

Acrocephalus newtoni 

Neomixis tenella 

Neomixis striatigula 

Nesillas typica 

Cisticola cherina 

Newtonia archboldi 

Newtonia brunneicauda 

Terpsi phone mutata (R) 

Nectarinia soui manga (R) 

Nectar inia notata (R) 

Lonchura nana (R) 

Zosterops maderaspatana 

Ploceus sakalava 

Foudia madagascariensis (R) 

Dicrurus forficatus (R) 



Mammals 

Lepilemur leucopus 
Microcebus murinus 
Lemur fulvus 
Pteropus rufus 
Setifer setosus 



Lemur catta 
Propithecus verreauxi 
Cheirogaleus medius 
Tenrec ecaudatus 
Eliurus sp 



Reptiles 

Acrantophis dumerilii 
Dromicodryas bernieri 



Leioheterodon madagascariensis 
Leioheterodon sp. 



Captive colonies of Geochelone radiata and Pyxis arachnoides. 

Information used in this account has been kindly provided by Sheila O'Connor and Mark 
Pidgeon. 



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VII. OTHER IMPORTANT AREAS 

The diversity and degree of endemism of the fauna and flora of Madagascar are such that 
almost all areas of undegraded natural vegetation on the island will be of considerable 
biological interest and value. Any selection of areas outside the Reserves Naturelles Integrates 
and the National Parks as sites of particular importance will thus to some extent be arbitrary, 
especially as knowledge of all these sites (as indeed of the protected areas themselves) is more 
or less incomplete. 

In terms of diversity of both animal and plant species, the most important areas are 
undoubtedly those with surviving low to mid-altitude rain forest; these are most extensive in 
the north-east of the island, from around 14°S to 18°S, although rain forest still persists along 
the length of the eastern escarpment to the far south. Identification of areas of particularly 
high diversity requires further study: Perrier de la Bathie noted (in La vegetation malgache 
1921, Marseille and Paris) that the whole of the eastern rain forest, to an altitude of ca 800 m, 
formed a single highly complex formation with essentially the same facies. Vegetatively the 
forest varied very little from north to south, but somewhat more so from east to west, this 
latter being almost certainly a reflection of increasing altitude. Floristic composition did 
appear to change somewhat, with gradual replacement of species by equivalent congeners, 
which however left the appearance and structure of the forest unchanged; the implication is 
that species diversity also remains relatively unchanged. Variations in faunistic composition 
are less clear; although, for example, more bird species appear to have been recorded in the 
northern half of the rain forest belt than in the southern half, this may well be a product of 
greater observer effort - the diverse avifaunas recorded recently at ca 21°S at Ranomafana 
(q.v.) and at ca 24°S in R.N.I. No. 11 (Andohahela) tend to bear this out. 

The following sites or regions have, however, had attention drawn to them in the literature or 
have emerged in the compilation of this report as areas of particular richness or interest; they 
should still be considered as examples or indicators. 



RAIN FOREST AREAS 



FORESTS OF MAROANTSETRA 



Forests in this area (the Antongil Bay region) include those of Ambohitsitondrona, Beanana and 
Ambohivoangy. Records cited from 'Ahitsitondrona' are thought to refer to 

Ambohitsitondrona. Strict protection of remaining forest areas here was advocated in the 
Proceedings of the 1970 Conference on Conservation in Madagascar. 

FAUNA 



Birds 

Ardeola idae (R) 
Lophotibis cristata 
Aviceda madagascariensis 
Eutriorchis astur 
Circus mail lard i (R) 
Accipiter madagascariensis 
Accipiter francesii (R) 
Falco zoniventris 
Mesitornis unicolor 
Canirallus kioloides 
Sarothrura insularis 
Actophilornis albinucha 
Glareola ocularis 



Alectroenas madagascariensis 
Coracopsis vasa (R) 
Coracopsis nigra (R) 
Agapornis cana 
Coua delalandei (Ex) 
Coua serriana 
Coua reynaudii 
Otus rutilus (R) 
Asio madagascariensis 
Caprimulgus enarratus 
Zoonavena grandidieri (R) 
Ipsidina madagascariensis 
Brachypteracias leptosomus 



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An environmental profile of Madagascar 



Brachypteracias squamiger 
Atelornis pittoides 
Leptosomus discolor (R) 
Philepitta castanea 
Neodrepanis coruscans 
Phedina borbonica (R) 
Coracina cinerea (R) 
Phyllastrephus madagascariensis 
Phyllastrephus zosterops 
Phyllastrephus tenebrosus 
Tylas eduardi 

Calicalicus madagascariensis 
Schelba rufa 
Vanga curvirostris 
Xenopirostris polleni 
Leptopterus madagascarinus (R) 
Oriolia bernieri 
Euryceros prevostii 
Hypositta corallirostris 



Copsychus albospecularis 
Monticola sharpei 
Neomixis tenella 
Neomixis viridis 
Neomixis striatigula 
Oxylabes madagascariensis 
Mystacornis crossleyi 
Acrocephalus newtoni 
Nesillas typica 
Rand i a pseudozosterops 
Newtonia amphichroa 
Newtonia brunneicauda 
Pseudobias wardi 
Terpsi phone mutata (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Lonchura nana (R) 
Foudia omissa 
Saroglossa aurata 



Mammals 

Propithecus diadema 
Hapalemur griseus 
Indri indri 
Varecia variegata 



Phaner furcifer 
Oryzorictes talpoides 
Microgale talazaci 



Reptiles 

Brookesia thieli 
*Typhlops ocularis 
Lycodryas betisleanus 



Chamaeleo linotus 
Geodipsas injralineata 



Nonmarine Molluscs 

Boucardicus beananae (Maroantsetra) 
*Boucardicus nanus (Ambohivoangy) 

Ampelila cerina (Maroantsetra) 

Ampelita fulgurata (Ambohitsitondrona) 

Ampelita lamarei (Ambohitsitondrona, Ambohivoangy) 

Ampelita lanx (Ambohivoangy) 

Ampelila stragulum (Maroantsetra, Ambohivoangy) 

Kalidos feneriffensis (Maroantsetra) 

Kalidos hestia ('Ahitsitondrona') 

Kalidos humbloti (Ambohivoangy) 

Kaliella ahitsitondronae ('Ahitsitondrona') 

Clavator moreleti (Maroantsetra) 

Helicophanta amphibulima (Maroantsetra) 

Fauxulus millotti (Ambohitsitondrona) 

Acroptychia aequivoca (Ambohivoangy) 
*Cyclotus millotti (Ambohivoangy) 

Macrochlamys stumpfi (Ambohivoangy) 

Tropidophora goudoliana (Ambohitsitondrona, Ambohivoangy, Beanana) 

Tropidophora perinelensis (Ambohivoangy) 

Tropidophora pulchella (Ambohitsitondrona) 

Tropidophora tricarinata (Ambohitsitondrona, Ambohivoangy, Beanana) 

Trochonanina millotti ('Ahitsitondrona') 
*Omphalotropis arbusculae (Ambohivoangy) 



•158- 



Other important areas 



MASQAT. A PENINSULA 



The Masoala Peninsula lies to the east of Antongil Bay and is contiguous with the preceding 
area. Reserve Naturelle Integrate No. 2 was situated in the north-eastern part of the peninsula, 
covering an area of 27 682 ha within the limits 15°17 , -15°24 , S and 50 o 13-50°30'E; it was 
de-gazetted in 1964. 

FAUNA 

Birds 

Most of the following species have been recorded by O. Langrand (in litt., 28.10.86). 



Lophotibis cristata 

Aviceda madagascariensis 

Eutriorchis astur 

Polyboroides radiatus 

Accipiter henstii 

Accipiter madagascariensis 

Accipiter f ranee sii (R) 

Buteo brachypterus 

Falco newtoni (R) 

Falco zoniventris 

Margaroperdix madagarensis 

Mesitornis unicolor 

Turnix nigricollis 

Dryolimnas cuvieri 

Canirallus kioloides 

Sarothrura insularis 

Streptopelia piclurata (R) 

Treron austral is (R) 

Alectroenas madagascariensis 

Coracopsis vasa (R) 

Coracopsis nigra (R) 

Agapornis cana 

Cuculus rochii (R) 

Coua serriana 

Coua reynaudii 

Coua cristata 

Coua caerulea 

Centropus toulou (R) 

Tylo soumagnei 

Otus rutilus (R) 

Asio madagascariensis 

Capri mulgus madagascariensis (R) 

Caprimulgus enarratus 

Zoonavena grandidieri (R) 

Alcedo vintsioides (R) 

Ipsidina madagascariensis 

Eurystomus glaucurus (R) 

Brachypteracias leptosomus 

Brachypteracias squamiger 

Atelornis pittoides 



Atelornis crossleyi 
Leptosomus discolor (R) 
Philepitta castanea 
Neodrepanis coruscans 
Phedina borbonica (R) 
Motacilla flaviventris 
Coracina cinerea (R) 
Phyllastrephus madagascariensis 
Phyllastrephus zosterops 
Hypsi petes madagascariensis (R) 
Calicalicus madagascariensis 
Schetba rufa 
Vanga curvirostris 
Xenopirostris polleni 
Leptopterus viridis 
Leptopterus chabert 
Leptopterus madagascarinus (R) 
Oriolia bernieri 
Euryceros prevostii 
Hypositta corallirostris 
Tylas eduardi 
Copsychus albospecularis 
Monticola sharpei 
Nesillas typica 
Newtonia brunneicauda 
Neomixis tenella 
Neomixis viridis 
Neomixis striatigula 
Pseudobias wardi 
Terpsiphone mutata (R) 
Oxylabes madagascariensis 
Mystacornis crossleyi 
Nectarinia soui manga (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Ploceus nelicourvi 
Foudia madagascariensis (R) 
Foudia omissa 
Lonchura nana (R) 
Saroglossa aurata 



159- 



An environmental profile of Madagascar 

'SIHANAKA FOREST' 

This is the name given by explorers to that part of the eastern rain forest, from the eastern 
coast to the Mangoro valley, east and south of Lake Alaotra, and in particular in the hinterland 
of Toamasina, especially between the towns of Didy and Fito. As discussed in the data sheet 
for Eutriorchis astur, (see Appendix 3. A), this name is technically a misnomer as the Sihanaka 
people live to the west of the rain forest proper, which is inhabited by the Betsimisaraka. As 
with the forests of Maroantsetra, this term cannot be applied precisely to a geographical entity, 
but rather to remaining areas of forest in the region. 

A number of threatened rain forest birds are best known from this region, and preservation of 
the remaining forest here is strongly advocated as a vital step in the conservation of these 
species. 

FAUNA 



Birds 

Lophotibis cristata 
A viced a madagascariensis 
Eutriorchis astur 
Accipiter henstii 
Falco zoniventris 
Mesitornis unicolor 
Sarothura insularis 
Gallinago macrodactyla 
Coracopsis nigra (R) 
Coua serriana 
Coua reynaudi 
Tyto soumagnei 
Otus rutilus (R) 
Asio madagascariensis 
Caprimulgus enarratus 
Ipsidina madagascariensis 
Brachypteracias leptosomus 
Brachypteracias squamiger 
Atelornis pittoides 
Atelornis crossleyi 
Leptosomus discolor (R) 
Philepitta castanea 
Neodrepanis coruscans 



Neodrepanis hypoxanlha 
Phyallastrephus zosterops 
Phyllastrephus tenebrosus 
Phyllastrephus cinereiceps 
Tylas eduardi 
Schetba rufa 
Xenopirostris polleni 
Oriola bernieri 
Hypositta corallirostris 
Copsychus albospecularis 
Monticola sharpei 
Neomixis tenella 
Neomixis viridis 
Hartertula flavoriridis 
Oxylabes madagascariensis 
Mystacomis crossleyi 
Crossleyia xanthophrys 
Dromaeocercus brunneus 
Newtonia amphichroa 
Newtonia fanovanae 
Pseudobias wardi 
Zosterops maderaspatana 



RANOMAFANA 

Ranomafana (21°16'S, 47°28'E) is situated ca 45 km north-east of Fianarantsoa in 
central-eastern Madagascar. A 1969 inventory of the forest domain of Madagascar quotes a 
forested area of 22 730 ha shared by the adjacent cantons of Ranomafana and Tsaratanana. 
This forest was not accorded any protected status at that time and the extent of surviving forest 
is unclear. This is the only site where Hapalemur simus is definitely known to survive, and the 
otherwise rarely recorded Lemur rubriventer has been described as 'abundant' here (P. Wright in 
litt. to S. O'Connor and M. Pidgeon, 22.08.86). Six threatened bird species have been recorded 
here since 1984 and for several Ranomafana marks a southern extension of the known range. 
N.B. There are at least two other localities named Ranomafana in eastern Madagascar. The 
following records were provided by O. Langrand and M. Nicoll {per O. Langrand, in litt., 
24.12.86). 



•160- 



FAUNA 



Other important areas 



Birds 

Aviceda madagascariensis 

Polyboroides radiatus 

Accipiter henstii 

Buteo brachypterus 

Falco newtoni (R) 

Mesitornis unicolor 

Dryolimnas cuvieri 

Canirallus kioloides 

Sarothrura insularis 

Streptopelia picturata (R) 

Treron austral is (R) 

Alectroenas madagascariensis 

Coracopsis vasa (R) 

Coracopsis nigra (R) 

Cuculus rochii (R) 

Coua reynaudii 

Coua caerulea 

Centropus toulou (R) 

Olus rutilus (R) 

Asio madagascariensis 

Caprimulgus madagascariensis (R) 

Caprimulgus enarratus 

Zoonavena grandidieri (R) 

Alcedo vintsioides (R) 

Ipsidina madagascariensis 

Eurystomus glaucurus (R) 

Brachypteracias leptosomus 

Atelornis pittoides 

Atelornis crossleyi 

Leptosomus discolor (R) 

Philepitta castanea 

Neodrepanis coruscans 

Phedina borbonica (R) 

Motacilla flaviventris 

Coracina cinerea (R) 

Phyllastrephus zosterops 



Phyllastrephus cinereiceps 
Phyllastrephus madagascariensis 
Hypsi petes madagascariensis (R) 
Calicalicus madagascariensis 
Schetba rufa 
Vanga curvirostris 
Xenopirostris polleni 
Leptopterus madagascarinus (R) 
Leptoplerus viridis 
Leptopterus chabert 
Tylas eduardi 
Copsychus albospecularis 
Monticola sharpei 
Acrocephalus newtoni 
Nesillas typica 
Cisticola cherina 
Dromaeocercus brunneus 
Dromaeocercus seebohmi 
Newtonia brunneicauda 
Neomixis tenella 
Neomixis viridis 
Neomixis striatigula 
Pseudobias wardi 
Terpsi phone mutata (R) 
Oxylabes madagascariensis 
Crossleyia xanthophrys 
Mystacornis crossleyi 
Nectar inia soui manga (R) 
Nectarinia notata (R) 
Zosterops maderaspatana 
Ploceus nelicourvi 
Foudia omissa 

Foudia madagascariensis (R) 
Lonchura nana (R) 
Dicrurus forficatus (R) 



Mammals 

Cheirogaleus major 
Microcebus rufus 
Avahi laniger 
Propithecus diadema 
Hapalemur simus 
Lemur fulvus 
Lemur rubriventer 
Lepilemur sp 
Fossa fossana 



Galidia elegans 
Galidictis fasciata 
Cryptoprocta ferox 
Hemicenteles semispinosus 
Setifer setosus 
Tenrec ecaudatus 
Microgale thomasi 
Nesomys rufus 
Brachytarsomys albicauda 



Amphibians 

Helerixalus alboguttatus 
Anodonthyla boulengeri 
Plethodontohyla notostica 



Mantidactylus liber 
Mantidactylus blommersae 
Boophis hillenii 



Reptiles 

Zonosaurus aenus 
Chamaeleo nasutus 
Phelsuma lineata 



Zonosaurus sp 
Chamaeleo brevicornis 
Sanzinia madagascariensis 



-161- 



An environmental profile of Madagascar 
NON RAIN FOREST AREAS 



ANKARANA MASSIF/AMBILOBE KARST 

This area, in the far north of the island (ca 13°S, 49°E) is sometimes known as the Ankara, 
under which it has been declared a special reserve of 18 220 ha, set up in 1956 (not to be 
confused with the Namoroka- Kelifely-Ankara karst in Mahajanga province, part of which is 
R.N.I. 8). 

The area, the northernmost sizeable karst area on the island, has the longest known cave system 
in Madagascar - the Grotte d'Andrafiabe, which has been extensively surveyed . 

FAUNA 



Birds 

Ardeola idae (R) 
Lophotibis cristata 
Anas bernieri 
Polyboroides r ad i at us (R) 
Streptopelia picturata (R) 

Mammals 

Lemur fulvus 
Fossa fossana 

Reptiles 

Homopholis boivini 
Lygodactylus rarus 
*Chamaeleo petteri 
Androngo all audi 
Ramphotyphlops braminus 
Liophidium there zieni 



Mesitornis variegata 
Actophilornis albinucha 
Pterocles personatus 
Nesillas typica 



Lemur coronatus 



*Lygodactylus expectatus 
*Phyllodactylus homalorhinus 
Amphiglossus waterloti 
Androngo elongatus 
Pseudoxyrhopus microps 



The area is also an important refuge for the Nile Crocodile Crocodylus niloticus. 

Nonmarine Crustacea 

Nine species of amphipods were collected in the area by the 1981 Southampton University 
Expedition, four of them new to science: 



Amphipoda 

Caridina parvocula sp nov. 
Caridina unca sp nov. 
Caridina nilotica 
Parisia dentata sp nov. 
Parisia microphthalma 

Nonmarine Molluscs 

Tropidophora cuvieriana 
Tropidophora deliciosa 
Tropidophora deshayesiana 
Tropidophora humberti 



Caridina crurispinata sp nov. 
Caridina norvestica 
Caridina isaloensis 
Parisia macrophthalma 



Tropidophora milloti 
Tropidophora surda siana 
Helicophanta socii 
Kalidos humbloti 



1 Southampton University Madagascar Expedition 1981 Final Report. Unpd, 136 pp. 
Radofilao, T. (1977). Ann. Univ. Madagascar Serie Sci, Nat. and Math, 14: 195-204. 



•162- 



Other important areas 



ANKARATRA MASSIF 



The Ankaratra is a volcanic massif situated about 70 km south of Antananarivo (see Part 
I.2.iv.). As early as 1950, much of the primary forest which originally covered the massif had 
already disappeared, and the rest was going fast. In view of the importance of this area, both 
botanically and zoologically, and the urgent need for a zoological inventory, efforts were being 
made between 1947 and 1958 by the staff of I.R.S.M. to collect material from the remaining 
wooded areas. 

Within the massif is situated the Manjakatompo forest station, which contains a vestige of 
natural high altitude forest, a pinetum, and clear streams (populated with introduced trout) and 
lakes. There are all-year tracks, one leading to the summit of Tsiafajavona, at 2643 m the 
highest point in the massif. 

FAUNA 



Birds 

Anas melleri 
Buteo brachypterus 
Margaroperdix madagarensis 
Leptosomus discolor (R) 
Mirafra hova 
Motacilla flaviventris 
Monticola sharpei 
Nesillas typica 



Dromaeocercus seebohmi 
Newtonia brunneicauda 
Terpsiphone mutata (R) 
Nectarinia souimanga (R) 
Zosterops maderaspatana 
Lonchura nana (R) 
Foudia madagascariensis (R) 
Foudia omissa 



Mammals 

Microgale dobsoni 



Amphibia (those marked (M) recorded at Manjakatompko) 



Plethodontohyla tuber at a (M) 
Tomopterna labrosa 
Mantidactylus domerguei (M) 
Boophis erythrodactylus (M) 
Boophis williamsi (M) 



*Pseudohemisus pustulosus 
Mantidactylus aerumnalis 
Mantidactylus pauliani (M) 
Boophis microtympanum 



Reptiles 

*Millotosaurus mirabilis 
Phyllodactylus homalorhinus 
Pseudoxyrhopus imerinae 



Phelsuma barbouri 
Mabuya madagascariensis 



Nonmarine molluscs (all from Manjakatompo) 
Clavator bathiei 
Sitala amabilis 
Sitala gaudens 
Clavator moreleti 



Macrochlamys stumpfii 
Vitrina madagascariensis 
Acroptychia aequivoca 



Crustacea (blind endemic species, recorded at Manjakatompo) 
Isopoda: 

Styloniscus albidus Suarezia differens 

Didima humilis Ankaratridium caecum 

Microcercus rotund if rons Armadillo ankaratrae 

Microcercus mascarenicus Bethalus carinatus 
Akermania sylvatica 

also recorded: 

Brycoyclops ankaratranus (forest of Ambahona) 
Astacoides madagascariensis caldwelli 



163- 



An environmental profile of Madagascar 

LAKE IHOTRY 

This lake, situated at 21°59'S, 43°36'E, just south of the Mangoky River, is cited as an area of 
considerable importance for waterbirds. Four threatened birds occur here, two, Tachybaptus 
pelzelnii and Ardea humbloti associated with the lake, and two, Monias benschi and Uratelornis 
chimaera, occurring in subdesert habitat adjacent to the lake. 

FAUNA 



Birds 

Tachybaptus pelzelnii 
Ardea humbloti 
Lophotibis cristata 
Anas bernieri 
Aviceda madagascariensis 
Polyboroides radiatus 
Circus maillardi (R) 
Accipiter henstii 
Accipiter madagascariensis 
Accipiter francesii (R) 
Buteo brachypterus 
Falco newtoni (R) 
Falco zoniventris 
Monias benschi 
Turnix nigricollis 
Actophilornis albinucha 
Pterocles personatus 
Streptopelia picturata (R) 
Treron australis (R) 
Coracopsis nigra (R) 
Coracopsis vasa (R) 
Agapornis cana 
Coua gigas 
Coua cursor 
Coua ruficeps 



Coua cristata 
Centropus toulou (R) 
Ninox superciliaris 
Caprimulgus madagascariensis (R) 
Ipsidina madagascariensis 
Uratelornis chimaera 
Leptosomus discolor (R) 
Phedina borbonica (R) 
Phyllastrephus madagascariensis 
Hypsipetes madagascariensis 
Calicalicus madagascariensis 
Xenopirostris xenopirostris 
Leptopterus chabert 
Copsychus albospecularis 
Neomixis tenella 
Neomixis striatigula 
Acrocephalus newtoni 
Thamnornis chloropetoides 
Newtonia brunneicauda 
Newtonia archboldi 
Terpsi phone mutata (R) 
Nectarinia souimanga (R) 
Zosterops maderaspatana 
Ploceus sakalava 
Dicrurus forficatus (R) 



ZOMBITSE FOREST 

The Zombitse Foret Classee covers 21 500 ha on gently undulating hills which form the 
western slopes of a sandstone dome. Soils are sandy with a very thin humus layer. There are 
no water courses in the forest although it acts as an important watershed, feeding springs that 
form tributaries of the Tehaza River which supplies extensive rice paddies 25 km south of 
Sakaraha. Climate is dry tropical with annual rainfall of ca 750 mm (maximum in December) 
and a dry season from May to October characterised by morning fog and heavy dew. The 
forest is the southernmost western domain dry deciduous forest. Canopy is dense, around 
15-20 m tall, and is dominated by Securinega seyrigii, Cedrelopsis grevii. Commiphora arofy, 
Khaya madagascariensis and Euphorbia anterophora. Herbaceous strata are very weakly 
developed. Illegal forestry was carried out from 1974 to 1981, using modern techniques and 
materials; all exploitable timber species were taken and Hazomalanga Hemandia voyroni 
(Hernandiaceae) has completely disappeared from the canopy. Exploitation has continued 
using traditional techniques to produce charcoal and building planks. Route Nationale 7 
crosses the forest and most disturbance appears to be concentrated here. 

The Zombitse Forest holds one of only two or three known tiny populations of Phyllastrephus 
apperti and Monticola bensoni has been recorded there in the non-breeding season. The 
gecko Phelsuma standingi is also confined to the region. 



-164- 



FAUNA 



Other important areas 



Birds 

Aviceda madagascariensis 

Accipiter francesii (R) 

Buteo brachypterus 

Falco newtoni (R) 

Falco zoniventris 

Margaroperdix madagarensis 

Turnix nigricollis 

Dryolimnas cuvieri 

Pterocles personatus 

Streptopelia picturata (R) 

Treron australis 

Coracopsis vasa (R) 

Coracopsis nigra (R) 

Agapornis cana 

Cuculus rochii 

Coua gig as 

Coua ruficeps 

Coua cristata 

Cent ro pus toulou (R) 

Otus rutilus (R) 

Ninox superciliaris 

Caprimulgus madagascariensis (R) 

Zoonavena grandidieri (R) 

Alcedo vintsioides (R) 

Eurystomus glaucurus (R) 

Leptosomus discolor (R) 



Mirafra hova 
Phedina borbonica (R) 
Motacilla flaviventris 
Coracina cinerea (R) 
Phyllastrephus madagascariensis 
Phyllastrephus apperti 
Hypsi petes madagascariensis (R) 
Calicalicus madagascariensis 
Vanga curvirostris 
Falculea palliata 
Leptopterus viridis 
Leptopterus chabert 
Leptopterus madagascarinus (R) 
Copsychus albospecularis 
Nesillas typica 
Cisticola cherina 
Newtonia brunneicauda 
Neomixis tenella 
Neomixis striatigula 
Terpsiphone mutata (R) 
Nectarinia souimanga (R) 
Zosterops maderaspatana 
Foudia madagascariensis (R) 
Saroglossa aurata 
Dicrurus forficatus (R) 



Mammals 

Microcebus murinus 
Phaner furcifer 
Propithecus verreauxi 
Lemur catta 
Lemur fulvus 
Lepilemur ruficaudatus 



Cryptoprocta ferox 
Tadarida sp. 
Tenrec ecaudatus 
Setifer setosus 
Echinops telfairi 
Geogale aurita 



Reptiles 

Phelsuma standingi 



The above information is taken from the report on the first phase of WWF 
programme in Madagascar, by M.E. Nicoll and O. Langrand. 



Protected areas 



165- 



APPENDIX 1. ENVIRONMENTAL LEGISLATION 



Legislation concerning the environment and natural resources both terrestrial and marine in 
Madagascar has been covered by the Rapport National pour Madagascar by Randrianarijaona 
and Razafimbelo (1983) prepared under the United Nations Environment Programme Regional 
Seas Programme; only a summary of such legislation will therefore by presented here. 



N.B. Protected area legislation is covered in detail in Part VI. 1. 
Part III.6. 



regulations covering forestry in 



A. SPECIES LEGISLATION. 



This is based primarily on the 1933 London Convention and on Ordonnance no. 60-126 
03.10.60. 



of 



The fauna has been divided into three categories (protected, game, vermin) and is now 
constrained under a series of dispositions controlling hunting and fishing. It is not clear if 
these have replaced the original tripartite classification or have merely amended it - there 
seems to have been no repeal of this earlier legislation. 

Some species can be captured for commercial ends, others only under 'exceptional 
circumstances'; such exploitation is controlled by Decrees (mainly no 61-093), which lay down 
the means, time and area of capture. Capture for scientific ends is generally authorized under 
the payment of a tax proportional to the scientific value of the species concerned (law 71-006). 

Some threatened species cannot be captured for any reason. In particular lemurs benefit from 
special restrictions on their being kept in captivity (Decree no 62-020). 



i. Protected Species 

All lemuriens 
Egretta gazetta dimorpha 
Bubuculus ibis ibis 
Phoenicopterus minor 
Testudo yniphora 
Acrantophis madagascariensis 
Typhleotris madagascariensis 

All newly introduced species, notably deer. 

ii. Vermin 

All Falco, Buteo, Astur 
Milvus migrans 
Foudia madagascariensis 
Ardea purpurea 
Coracopsis 
'Felis ocreata' 
Galidictis vitata 
Potamochaerus larvatus 



Dugong dugon 

Egretta alba melanorhynchus 
Phoenicopterus ruber 
Lophotibis cristata 
Testudo radiata 
Acrantophis dumerili 
Typhleotris pauliani 



Milvus parasitus 
Ardea cinerea 
Corvus albus 
Crocodylus niloticus 
Galidictis striata 
Viverricula schlegeli 
Pteropus rufus 



N.B. Nomenclature follows that given in relevant ordonnances 



-167- 



An environmental profile of Madagascar 

B. INTERNATIONAL AGREEMENTS 

Madagascar has affiliated to the following international treaties and conventions, concerned 
with the environment: 

i. World Heritage Convention 6 Feb 1983 (Ratified) 

[As of 1986, no Madagascan sites had been formally proposed for inclusion as World 
Heritage Sites.] 



n. 



in. 



Bonn Convention on Migratory Species 
1933 African Convention 



Signed but not ratified 
9 Oct 1969 (Ratified) 



iv. 1973 Convention on International Trade in Endangered Species of Wild Fauna and Flora 
(CITES) 18 Nov 1975 (Ratified) 

The following species and groups of species which occur in Madagascar are listed in the 
Appendices to the CITES conventions. Trade in Appendix 1 species (or their products) is 
subject to strict regulation by ratifying nations, with trade for primarily commercial 
purposes banned; trade in Appendix 2 species is subject to monitoring by ratifying 
nations. 



APPENDIX I 



APPENDIX II 



MAMMALIA 



All Lemurs 
Dugong dugon 



Cryptoprocta ferox 
Eupleres goudotii 
Eupleres major 
Fossa fossa 



AVES 



Tyto soumagnei 



Anas bernieri 

All Falconiformes 

All Psittaciformes 

Strigiformes (except that on Ap I) 



REPTILIA 



Geochelone radiata 
Ceochelone yniphora 
All Cheloniidae 
Crocodylus niloticus 
Acrantophis spp 
Sanzinia madagascariensis 



Testudinidae (except those on Ap I) 



Phelsuma spp 
Chamaeleo spp 



PLANTS 



Pachypodium spp. 
Cactaceae 
Cyatheaceae 
Cycadaceae 
Didieraceae 
Euphorbia spp 
Aloe spp 
Orchidaceae 



-168- 



APPENDIX 2. FAUNAL LISTS 

Annotated lists of species for the following groups are provided: 

Birds, divided into endemics (country and regional) and non-endemics; 

Mammals (all native non-marine species); 

Reptiles (all native species); 

Amphibians (all native species); 

Fish (endemic species and subspecies); 

Lepidoptera (Rhopalocera except family Hesperidae); 

Molluscs (freshwater and endemic terrestrial species); 

Nonmarine Crustacea (selected groups); 

These lists serve (together with data sheets for individual species in Appendix 3) as adjuncts to 
Part V, where the references used in their compilation can be found. 

Slightly different formats have been adopted for each group, though this, and any 
abbreviations used, are explained at the beginning of each list. 



-169- 



An environmental profile of Madagascar 

MADAGASCAR BIRDS 

1. ENDEMIC 

(R) indicates endemic to Madagascan region; all others are 
endemic to Madagascar. Threatened species are treated in 
detail in Appendix 3. 

Family PODICIPEDIDAE 

Tachvbaptus pelzelnii Madagascar Little Grebe 

INSUFFICIENTLY KNOWN: Common and widepread in 
recent past, suffered considerable decline around 
Antananariavo, Lake Alaotra, Lake Ihotry. Widespread but 
no longer common anywhere. Sealevel - 1800 m; absent only 
from subdesert in south. 

Tachvbaptus rufolavatus Alaotra Grebe 

ENDANGERED: Known chiefly from Lake Alaotra in 
north-east; also from west, although validity of many 
records away from L. Alaotra is questionable. Is in 
irreversible process of disappearing through hybridisation 
with T. ruficollis . 

Family ARDEIDAE 

Ardeola idae (R) Madagascar Pond-heron 

Occurs throughout; generally considered commoner in west 
than east. Winters in East and Central Africa; massive 
recent decline in only well known breeding area (ca 1500 
birds in 1945, 50 in 1974) may also have occurred elsewhere. 
Also on Seychelles. 

Ardea humbloti Madagascar Heron 

INDETERMINATE: Very thin distribution in west, chiefly 
in coastal and adjacent areas; rarely in east. Only 4 
breeding sites found: north, central west, south-west, off 
south-west coast. 

Family THRESKIORNITHIDAE 

Lophotibis cristata Madagascar Crested Ibis 

Widespread species; western subspecies widespread and 
common, eastern subspecies shows adaptability: recorded 
from secondary forest and plantations. Recorded from at 
least six protected areas. 



Family ANATIDAE 
Anas bernieri 



Madagascar Teal 



VULNERABLE: Little known, much persecuted, known 
from few sites on west coast only, total numbers probably 
very low. 



Anas melleri 



Meller's Duck 



Most frequently recorded east (from Vondrozo north to 
Andapa and on central plateau), though also recorded in 
west. Often hunted. Common in some localities but has 
suffered an apparent marked decline recently. Introduced 
Reunion and Mauritius. 

Aythya innotata Madagascar Pochard 

ENDANGERED: Poorly known, confined to lakes of 
north-central plateau, especially Lake Alaotra (but no 
records from there since 1930s). Also Lake Itasy, Lake 
Ambohibao, near Antsirabe, Ambatomainty, and Betsileo 
country. 

Family ACCIPITRIDAE 

Aviceda madagascariensis Madagascar Cuckoo-falcon 

Widely distributed in a variety of habitats; uncommon 
though probably overlooked. Nestlings eaten and adults 
snared at nest. 

Haliaeetus vociferoides Madagascar Fish-eagle 

ENDANGERED: Central west coast to North. Main areas: 
between Antseranana and Nosy Be, Lake Kinkony, 
Antsalova, coastline between Mangoky and Fierenana 
rivers. Population most recently estimated at around 30 
pairs. 



Eutriorchis astur Madagascar Serpent Eagle 

ENDANGERED: Very poorly known, last seen by 
ornithologist 50 years ago. Humid east, dense rain forest. 
Ampa Simanavy (in Mangoro Valley), Rogez, Maroantsetra, 
Farafangana, Marojejy Reserve (possible records only). 
Extremely rare. 

Polvboroides radiatus Madagascar Harrier-hawk 

Widely distributed in variety of wooded habitats and open 
country. Fairly common except centre, extreme north, 
south, and south-west. Adapted to degraded/secondary 
woodland. 

Circus maillardi (R) Madagascar Harrier 

Rare; marshes and grassland of east and west; reported as 
locally fairly common (Lake Ihotry, near Vohimarina). Also 
Reunion and Anjouan. 

Accipiter hensti Henst's Goshawk 

Widely distributed in variety of habitats but uncommon and 
in decline. Treated as harmful species under government 
decree, still in force. 

Accipiter madagascariensis Madagascar Sparrowhawk 
Largely confined to areas below 1000 m; extremely rare in 
east, uncommon in west, somewhat commoner in subdesert 
south-west. Forest, open woodland, wooded savanna. 

Accipiter francesii (R) Frances's Sparrowhawk 

Common in dry and humid and even degraded forest 
throughout. Most common in north-east. Also on 
Comoros. On government list of harmful species. 0-1800 m. 

Buteo brachypterus Madagascar Buzzard 

Common throughout, most records from east, locally 
abundant. Prefers savanna, forest edges, adapted to 
exploited/degraded forest. 0-1800 m. 

Family FALCONIDAE 
Falco newtoni (R) Madagascar Kestrel 

Reasonably common throughout especially central plateau; 
prefers open areas to dense forest; found in degraded habitat 
and cultivated areas. Commonest raptor. Also on Aldabra. 
On government list of harmful species. 0-2000 m. 

Falco zoniventris Banded Kestrel 

Widely distributed but rare, may be commonest in 
north-east and south-west. Probably overlooked. Treated 
as harmful species under government decree, still in force. 
Forest, wooded savanna, cultivated areas. Insectivorous. 

Family PHASIANIDAE 

Margaroperdix madagarensis Madagascar Partridge 
Widely distributed except dense forest, south-west and 
south. Up to 2500 m in Tsaratanana Massif. Rice, savanna, 
afforestation areas, secondary woodland. Common but 
diminishing through hunting. Introduced Reunion and 
Mauritius (now extinct on latter). 

Family MESITORNITHIDAE 
Mesitornis variegata White-breasted Mesite 

RARE: Terrestrial forest bird, currently known from only 
two sites: Ankarafantsika, which is, however a protected 
area, and north-east of Morondanva. In past known from 



near Tsarakibany and perhaps 
Haut-Sambirano. 



the Analalava and 



Mesitornis unicolor Brown Mesite 

INSUFFICIENTLY KNOWN: Rainforest, perhaps wider | 
distribution than known, possibly throughout East. Most 
reliable records from circle with diameter Antananarivo to 
Toamasina and from south-east corner. 

Monias benschi Subdesert Mesite 

RARE: Terrestrial, restricted range in subdesert 

(south-west) between Mangoky and Fierenana rivers. Sea < 

level to 130 m. Common over much of range, at times 

abundant. 



■170- 



Grey-throated Rail 



Family RALLIDAE 
Mentocrex kioloides 
(= Canirallus kioloides ! 
East from Manombo north to the north-west (Sambirano); 
reaches central high plateau (1450 m). Common in humid 
forest (1 pair per km: Perinet). Can adapt to secondary 
woodland. 

Sarothrura insularis Madagascar Flufftail 

East, north-west (Sambirano), north (Mt. d'Ambre). 
Secondary brush/grassland on forest edge. 0-2300 m. 
Commonest in East. 

Sarothrura watersi Slender-billed Flufftail 

INSUFFICIENTLY KNOWN: Recorded from 3 widely 
separated areas: south-east Betsileo (south-centre), Andapa 
(north-east), Antananarivo. Small swamps (association with 
Cvjjerus). 

Dryolimnas cuvieri White-throated Rail 

Rather common throughout in forests. 0-1800 m, but rare 
above 1100 m. Edges of marshes and streams. Commonest 
north-west and humid eastern forest. Rarer on central 
plateau. 



Rallu8 madagascarienesis Madagascar Rail 

East, north-east (Andapa) and central plateau. 
Commoner at higher altitudes. Few records. 



0-1800 m. 



Amaurornis olivieri Sakalava Rail 

INSUFFICIENTLY KNOWN: Rare and localised; known 
from only three widely separated areas in the Sakalava 
country in west. Recorded from Antsalova, Ambaratabe, 
Tsiribahina River, Nosy Ambositra. Marshes and streams. 

Family TURNICIDAE 

Turnix nigricollis Madagascar Buttonquail 

Widespread and common in variety of open habitats, 
especially high plateau. 0-1900 m. Introduced to Mauritius 
and Reunion. 

Family JACANIDAE 

Actophilornis albinucha Madagascar Jacana 

Centre-east (from Mahanoro north); north-east, north-west, 
west down to Tulear; north-central plateau. Commonest in 
west, north-west and north-east; less recorded from forested 
east, where it occurs in forest clearings and rice-cultivation 
areas. 

Family GLAREOLIDAE 

Glareola ocularis Madagascar Pratincole 

East from Fianarantsoa to north-east, north-west, northern 
savanna and west (Bekipay and Mangoky). Migratory to 
Africa. Rocks on perennial rivers, beaches, river edges. 

Family CHARADRIIDAE 
Charadrius thoracicus Madagascar Plover 

RARE: Restricted to coastal grassy areas of south-west 
though first described in east. Population probably under 
1000. 

Family SCOLOPACIDAE 

Gallinago macrodactvla Madagascar Snipe 

East, central plateau (as far west as Sakay), north 
(Tsaratanana) up to 2700 m. Common (2 pairs per hectare 
at Tampoketsa d'Ankazobe) but status needs watching since 
habitat disturbed by rice growing. 

Family PTEROCLIDIDAE 

Pterocle8 personatus Madagascar Sandgrouse 

West: common from Taolanaro (south-east) to north-west. 
Extends beyond Sambirano (between Amabanja and 
Antseranana). Sandy dry plains, savanna, sparsely wooded 
regions. 



Appendix 2: fauna! lists 

Family COLUMBIDAE 

Alectroenas madagascariensis Madagascar Blue Pigeon 
Most records from north (Sambirano, Tsaratanana, Andapa 
and Mt. d'Ambre). Also from length of the east. Common 
in dense forests, commoner in east than west where it 
migrates March-July. Numbers at Perinet reduced by 
hunting. 

Streptopelia picturata (R) Madagascar Turtle Dove 
Throughout east, west, subdesert (south-west), common in 
forest, brushland, plantations; less so in open areas. 
0-2000 m. Also on Comoros, Aldabara, Chagos. Introduced 
Seychelles, Mauritius, Reunion, Amirantes. 

Treron australis (R) Madagascar Green Pigeon 

Very common in wooded areas, 0-1000 m. Absent from high 
plateau. Hunted. Also Comoros. 

Family PSITTACIDAE 

Agapornis cana Grey-headed Lovebird 

Very wide distribution in a variety of habitats. Common 
throughout, but particularly in lower and less arid areas. 
Uses cultivated areas, rice fields, degraded forest. 

Coracopsis vasa (R) Greater Vasa Parrot 

Widespread. Woodland east, west, south, including 
degraded. Prefers coastal plains 0-1000 m. Visits cultivated 
areas. Common. Also Anjouan, Reunion. On government 
list of harmful species. 

Coracopsis nigra (R) Lesser Vasa Parrot 

Common throughout; woodland (including degraded) 
preferring denser forest and brush than C. vasa , thus less 
common in west than C. vasa . 0-2050 m. Also Comoros, 
Seychelles. On government list of harmful species. 



Family CUCULIDAE 
Coua caerulea 



Blue Coua 



East from Taolanaro to Sambirano (north-west) with relict 
population in dry wood at Bora (Antsohihy). Rain forest 
but also mangroves. 0-1800 m. 



Coua cristata 



Crested Coua 



Wide distribution in variety of habitats; predominance of 
records from west and centre, less from humid east where its 
presence marks a south- ward extension coinciding with rain 
forest destruction. 

Coua verreauxi Verreaux's Coua 

Highly restricted, only in dry south-west, between Fierenana 
and Menarandra rivers, but fairly common; occurs within 
Lake Tsimanampetsotsa Nature Reserve. Confined to area 
of thick coastal scrub on coral rag. 

Coua reynaudii Reynaud's Coua 

Common in eastern rainforest from Manombo north to 
north-east and Sambirano as far west as Mandraka in 
centre. Relict population in dry forest at Bora (Antsohihy). 
Up to 2500 m. 

Coua serriana Red-breasted Coua 

Northern part of humid east (one old record from 

south-east) from Sihanaka to Sambava. Common in its 
limited range. 0-1000 m. 

Coua ruficeps Red-capped Coua 

West from Faux Cap north to Mampikony in 2 discrete 
populations; south-west and west around Mahajanga. Less 
common in extreme south. Adaptable to degraded regions. 
Also in dry forest, savanna, sandy areas. 



Coua cursor Running Coua 

West from Morondava south to Cap Ste. Marie, 
brush, dry forest. 0-160 m. 



Desert 



Coua coquereli Coquerel's Coua 

West from Sakaraha north to Maromandia. Incursions into 
south-west sub-desert and Sambirano. Rarely in degraded 
forest. Common but local. 



•171- 



An environmental profile of Madagascar 



Coua gigas Giant Coua 

West: Toliara north to Mampikony. Recent records extend 
range to Berenty and Andohahela in south-east, west of 
Taolanaro. 0-700 m. Dry forest areas. Primary habitat 
only, not in degraded zones. Rather common in some areas. 



Coua delalandei Snail-eating Coua 

EXTINCT: Known chiefly from Nosy Borah 

Sainte-Marie) (north-east); no precise records 
mainland. Forest dweller, subsisting on molluscs. 



(lie 
from 



Cuculus rochii Madagascar Little Cuckoo 

Widespread and quite common, only absent from driest part 
of south; wooded areas savanna and marshes, 0-1800 m. 
Migrates from east to west (and on to Africa) during rainy 
season. 

Centropus toulou (R) Madagascar Coucal 

Common and widespread in wooded and brush areas 
throughout except for treeless central plateau regions where 
less recorded (though common at Sahavondronina). 
0-1800 m. Prefers less dense wooded areas; also in 
marshland and degraded woodland. Also on Mayotte, 
Aldabra. 

Family TYTONIDAE 

Tvto soumagnei Madagascar Red Owl 

INDETERMINATE: Known with certainty from humid 
rainforest only in centre-East; seen only once in past 50 
years. Recorded from circle whose diameter runs between 
Toamasina and Antananarivo. 

Family STRIGIDAE 
Otus rutilus (R) Madagascar Scops Owl 

Reported as common, scattered records from a wide area in 
forest/brushlands including partially exploited forest. Also 
on Comoros. 0-1800 m. 

Ninox superciliaris White-browed Owl 

All records from west and south-west between Ampotaka 
and Morondava except one from Marojejy. Not rare, forest 
and wooded savanna. 

Asio madagascariensis Madagascar Long-eared Owl 

Mostly in centre-east forest, as far west as Antananarivo. 
Also recorded west savanna (Tabiky) and Maromandia. 
0-1800 m. Rather rare but probably overlooked. 

Family CAPRIMULGIDAE 

Caprimulgus enarratus Collared Nightjar 

All records except two (Taolanaro and unknown locality in 
south-east) from centre-east and north-west (Sambirano) 
from dense evergreen forest. 0-1800 m. Quite common. 

Caprimulgus madagascariensis (R) Madagascar Nightjar 
East from Ambodiasy north to Antseranana. Records also 
from Antananarivo area, Mampikony and Toliara, Sakaraha, 
Lac Ihotry (west). Can adapt to degraded areas; forest and 
open brush. Also on Aldabra. 

Family APODIDAE 
Collocalia francica (R) Mascarene Swiftet 

Rare; East coast. Also on Mauritius and Reunion. 

Zoonavena grandidieri (R) Madagascar Spinetail 
Widely distributed forest dweller. 0-1000 m, common 
especially at lower altitudes. South-east, north-east, weBt, 
south-west; commonest in east. Also on Grand Comoro. 

Family ALCEDINIDAE 

Alcedo vintsioides (R) Madagascar Malachite 

(= Corythornis vintsioides ) Kingfisher 

Common east and west (and recorded from subdesert). No 
strict association with water. 0-1800 m. Found in degraded 
woodland, afforestations and on tideline. Also on Comoros. 



Ipsidina madagascariensis Red and White Kingfisher 

(= Cevx madagascariensis ) 

Widely distributed except extreme south-west; largest 
concentrations of records in centre. Quite common in east, 
rarer in west. Common on Mt. d'Ambre and Sambirano. 
0-1800 m. 

Family LEPTOSOMATIDAE 
Leptosomus discolor (R) Courol 

Widely distributed. Common in east in forests and 
secondary brush; in west and subdesert only found in denser 
savanna and wooded plains. 0-2000 m. Uses degraded areas 
of forest. May migrate to west during humid season. Also 
Mayotte, Anjouan. 

Family BRACHYPTERACIDAE 
Eurystomus glaucurus (R) Broad-billed Roller 
Found throughout, September to April, migrates to Africa 
for remainder of year. Plentiful even in east. Also on 
Comoros, Aldabra, vagrant to Reunion and Rodrigues. 

Brachypteracias leptosomus Short-legged Ground-roller 
RARE: Occurs in 2 discrete areas: north-east (Marojejy to 
around Maroantsetra) and centre-east (chiefly Sihanaka 
forest). Deep rainforest. Like all Ground-rollers probably 
overlooked. 

Brachypteracias squamigera Scaly Ground-roller 
RARE: Deep rainforest in centre and north-east: Marojejy, 
Andapa, Maroantsetra, Masoala, Soamiana, Sihanaka forest, 
Perinet, Rogez and perhaps in South-east. 0-1800 m. 

Atelornis pittoides Blue-headed Ground-roller 

Formerly regarded as rare, now known to be widespread 
from Mt. d'Ambre in north to Chaines Anosyennes in south 
and as far west as Mandraka. Occurs in several reserves. 
Dense evergreen forest. 

Atelornis crosslevi Rufous-headed Ground-roller 

RARE: Confined to deep rainforest in east, with records 
scattered from Tsaratanana Massif south to Vondrozo 
region. Rarest and least known Ground-roller. 

U ratelornis chimaera Long-tailed Ground-roller 

RARE: Restricted range within subdesert (South-west) 
between Mangoky and Fierenana rivers, ranging up to 80 m; 
distribution coincides with that of Didierea woodland, within 
which it may be locally abundant. 



Family PHILEPITTIDAE 
Philepitta castanea 
Common throughout east 
north-west (Sambirano). 
Mandraka where not rare. 



Velvet Asity 

from Taolanaro north to 

As far west as heights of 

0-1800 m. Can adapt to 



secondary/exploited woodland. 

Philepitta schlegeli Schlegel's Asity 

Confined to west (from Tsiandro north) and Sambirano; old 

north-east record. Dense forest, localised, but common in 

Sambirano. 

Neodrepanis coruscans Sunbind Asity 

East from Vondrozo north to Tsaratanana. Locally common 
in many places. 0-1800 m. Favours higher altitudes. Moves 
out into dense secondary brush. 

Neodrepanis hypoxantha Yellow-bellied Sunbird Asity 

INDETERMINATE: East, centre in forests east and 
perhaps south of Antananarivo and the Sihanaka forest. 
Known only from 13 specimens collected before 1930, and 
records of breeding in 1973 and 1976. 

Family ALAUDIDAE 

Mirafra hova Madagascar Bush Lark 

Widespread and common especially south-west and centre 
uplands in open ground, into forest clearings, degraded 
forest. 0-2500 m. 



-172- 



Appendix 2: faunal lists 



Family HIRUNDINIDAE 

Phedina borbonica (R) Mascarene Martin 

Fairly common but irregular distribution throughout east, 
west, subdesert. Breeding up to 2200 m (Tsaratanana). 
Forest, including degraded and afforestation areas. Also 
Reunion, Mauritius, vagrant to Seychelles. Migratory north 
to south. 

Family MOTACILLIDAE 

Motacilla flaviventris Madagascar Wagtail 

Widespread, except for south and south-west. Especially 
common on high plateau up to 2600 m. Cultivated ground, 
exploited forest, paddy fields. Possibly seasonal migrant. 

Family CAMPEPHAGIDAE 
Coracina cinerea (R) Madagascar Cuckoo-shrike 

Common throughout in a variety of habitats, including 
degraded areas, up to 2300 m. Occurs in mixed species 
foraging groups. Also on Comoros. 

Family PYCNONOTIDAE 

Hypsipetes madagascariensis (R)Madagascar Bulbul 
One of commonest birds, found in forest and brush 
throughout. 0-2500 m. Uses degraded habitat, afforestation 
areas. Also on Mayotte, Anjouan, Moheli, Aldabra, 
Seychelles. 

Phvllastrephus cinereiceps Grey-crowned Greenbul 
RARE: Possibly occurs throughout rainforests of east but 
known from only a few scattered sites. Inhabits ground 
cover of deep rainforest. Recorded from Fianarantsoa, 
Sihanaka forest, Fanovana, Ranomafana. 

Phvllastrephus tenebrosus Dusky Greenbul 
RARE: Recorded from only 3 areas in the east (Sihanaka 
forest, Pe>inet-Analamazaotra and Maroantsetra). May be 
overlooked. Probably confined to primary rainforest. 

Phvllastrephus zosterops Short-billed Greenbul 

East from near Vondrozo north to Mt. d'Ambre (including 
Tsaratanana). Dense humid forest plus secondary 

woodland. Variable in abundance throughout range. 

Phyllastrephus apperti Appert's Greenbul 

RARE: Known from only one locality (Zombitsy forest 
south-east of Ankazoabo) in south-west. Local and sparse 
within known range. Dense dry forest. 

Phyllastrephus madagascariensis Long-billed Greenbul 
Widespread: east from Taolanaro to Antalaha; west from 
Vohemar to Toliara (including Sambirano). Common. Uses 
secondary woodland. Absent from central massif. 

Family VANGIDAE 

Calicalicus madagascariensis Red-tailed Vanga 
East, Tsaratanana, north-west (Sambirano), south-west: 
common (less so in west); forest and woodland including 
partially exploited. Occurs in mixed species foraging 
groups. 0-2050 m. 



Schetba rufa 



Rufous Vanga 



East (from Vondrozo north to Andapa), west (from 
Sakaraha to Mahajanga area). Forest bird, locally common 
but absent from many places. Greatest density in primary 
forests. 

Vanga curvirostris Hook-billed Vanga 

Widespread and fairly common throughout east and west in 
wooded areas/forest into degraded/secondary growth, 
mangroves in north-west, plantations. Occurs in mixed 
species foraging groups. 

Xenopirostris xenopirostris Lefresnaye's Vanga 
Restricted to south-west subdesert where not uncommon in 
undergrowth of limestone collines on coast, especially with 
Didierea bush and Euphorbia . 



Xenopirostris damii Van Dam's Vanga 

RARE: Known this century only from Ankarafantsika 
(south-east of Mahajanga) which is however a protected 
area and where it occurs in fairly good numbers. Previously 
collected in north-west. Primary deciduous forest. 

Xenopirostris polleni Pollen's Vanga 

INDETERMINATE: Wide variety of localities in east 
primary rainforests (although type is from north-west 
coast). Suspected sight record from Marojejy; all other 
records from central part of east. Wider range than often 
stated. 

Falculea palliata Sicklebill 

Primarily west distribution, but including Berenty (near 
Taolanaro in South-east), Mt. d'Ambre and Sambava in 
north-east. Largest concentration of records is from west 
and south-west. Common, occurs in mixed species foraging 
flocks. 0-900 m. 

Leptopterus viridis White-headed Vanga 

(= Artamella viridis ) 

Widespread in a variety of habitats: woodland, primary 

humid and dry forest, coastal mangroves, brush, subdesert. 

Common. 

Leptopterus chabert Chabert's Vanga 

Widespread throughout east and west (except extreme north 
and Sambirano). Occurs in secondary/exploited forest, rice 
paddies and mangroves. 

Leptopterus madagascarinus (R) Blue Vanga 

(= Cvanolanius madagascarinus ) 

Common in forest and brush in east, Mt d'Ambre, 

Sambirano; less common in west, rarer in south ranging into 

edge of subdesert. Uses degraded habitat. 0-1800 m. Occurs 

in mixed species foraging groups. Also on Comoros 

(Moheli). Perhaps locally migratory. 



Oriolia bernieri 



Bernier's Vanga 



Treated as Indeterminate in previous Red Data Book but 
widely distributed in rain-forests. Probably overlooked in 
tree tops. Occurs in mixed species foraging groups. 
500-1000 m. 

Eurvceros prevostii Helmet Bird 

Local, total population may be low: primary forest in humid 

east from Fanovana to Andapa; never in degraded areas. 

Tvlas eduardi Tylas Vanga 

Occurs in the east from Vondrozo north to Tsaratanana 
(west to Andrangolsaka). Discrete western population found 
around Morondava south to south-east of Ankazoabe; rare 
in west, perhaps a mangrove dweller. In east in dense 
forest. Nowhere common. 

Hypositta corallirostris Coral-billed Nuthatch-Vanga 

Occurs in the east from Taolanaro north to Marojejy. Not 
rare, evergreen rainforest, near sea level - 1800 m. Possibly 
commoner at higher altitudes. 

Family MUSCICAPIDAE 
Copsvchus albospecularis Madagascar Magpie Robin 

Common throughout, especially in wooded areas/forest 
edges, neighbouring brush, even in interior of evergreen 
forest, and exploited/degraded areas. 

Monticola sharpei Forest Rockthrush 

(= P8eudocossyphus sharpei ) 

Found in the east from Ivohibe north to Mt. d'Ambre, west 

to Massif d'ltremo and Tsaratanana. Locally common: dense 

forest including secondary woodland, also high plateau 

grassland. 100-2200 m. 

Monticola imerina Littoral Rockthrush 

(= Pseudocossyphus imerina ) 

Local south-west (Mangoky) to Taolanaro (south-east). 

Common. Dunes, dry grassland with Euphorbia and 

Didierea . not in forest. 



■173- 



An environmental profile of Madagascar 



Monticola bensoni Benson's Rockthrush 

(= Pseudocos8yphus bensonO 

INSUFFICIENTLY KNOWN: probably quite widespread 

but as yet known only from a few dry rocky areas in 

south-west: Mangoky river region and north of Isalo massif. 

Keeps mostly to rocks/cliff faces. 

Neomixis striatigula Stripe-throated Jery 

East (north to Andapa) and South-west. Two subspecies: 
forest from Fanovana north to Andapa in upland areas, 
800-1800 m. More common southern subspecies extends as 
far east as Fianarantsoa. 

Neomixis viridis Green Jery 

East from Ivohibe north to Tsaratanana. All records from 
inland areas 1000-2050 m. Often in damp, mossy forest. 
Not very common. 



Neomixis tenella 



Common Jery 
Records from all areas 



Widespread and common. Records from all areas (few in 
central plateau) and variety of habitats, wooded subdesert, 
rainforest, severely degraded secondary woodland. 0-1200 m. 

Hartertula flavoriridis Wedge-tailed Jery 

Uncommon, east rainforests, stronghold Sihanaka forest; 
other records Vondrozo, Lakata (presumably Lakato), 
Perinet-Analamazaotra Special Reserve, Bejofo-Bealanan, 
Tsaratanana Nature Reserve. 

Oxvlabes madagascariensis Yellow-browed Oxylabes 
Rather common east, north-east, north (Mt. d'Ambre) from 
Manombo (with old record from Taolanaro). Most records 
from within circle whose diameter is 

Antananarivo-Toamasina. Dense rainforest. 0-1800 m. 

Mystacornis crossleyi Crossley's Babbler 

East from Manombo north to Andapa. Common, especially 

away from coast and at higher altitudes in dense humid 

forest. 0-1800 m. Occurs in foraging parties of mixed 

species. 

Acrocephalus newtoni Madagascar Swamp Warbler 

Widespread and rather common east, north-west, west, 
central massif; marshes, mangroves, rivers. 0-1800 m. 

Crossleyia xanthophrys Madagascar Yellowbrow 

INDETERMINATE: Confined to rainforest in centre-east 
with one record from north (Tsaratanana). Seen only twice 
in past 50 years but possibly overlooked. 

Nesillas typica Madagascar Brush Warbler 

Widespread and common; absent only from extreme 
South-east. Most records are from East, North, 

North-west. Variety of habitats; primary forest, brush, 
severely degraded secondary forest. 0-2750 m. 

Thamnornis chloropetoides Kiritika 

Restricted to south-west subdesert but common. Cap 
Sainte-Marie north to Lake Ihotry. Shrub, brush, small 
trees, often Euphorbia and Didierea in sandy, arid area. 

Cisticola cherina Madagascar Cisticola 

Common and widespread through variety of habitats. Most 
records are from east between Faroany river and Mohambo 
and from central massif. 0-2000 m. Savanna, partially 
exploited primary forest, swamp, cultivated areas, shoreline. 

Dromaeocercus seebohmi Grey Emu-tail 

Humid east from Ankaratra to Tsaratanana at altitude, 

900-2600 m. Common. Bushes, marshes, rivers. 

Dromaeocercus brunneus Brown Emu-tail 

Centre-east rainforest. Fairly common in 

Perinet-Analamazaotra Special Reserve; locally abundant in 
Sihanaka forest, Fierenana forest and even partially 
exploited forest at Nangarana. 



Randia pseudozosterops Rand's Warbler 

Rainforest. Originally considered very rare, now known to 

be commoner and fairly widespread. Also occurs in 

secondary woodland. Occurs in mixed species foraging 

groups. 

Newtonia brunneicauda Common Newtonia 

Widespread and common. Occurs in east from Manombo to 
Mt. d'Ambre and in west down to Ampotaka (south-west). 
Uses degraded woodland. 0-2000 m. 

Newtonia amphichroa Dark Newtonia 

Humid east from Ivohibe north to Mt. d'Ambre (two 
specimens from south-west were perhaps wrongly labelled). 
Not common. Dense forest. 500-1800 m. 

Newtonia archboldi Archbold's Newtonia 

Restricted to south-west and not common; Ampotaka north 
to Lake Ihotry in brush, low forest; often found in Euphorbia 
and Didierea brush. 

Newtonia fanovanae Red-tailed Newtonia 

INDETERMINATE: Known only from single specimen from 
humid forest, now cleared, in east-centre. If it is not an 
invalid taxon based on an aberrant bird, it is either 
overlooked, genuinely rare or extinct. 

Pseudobias wardi Ward's Flycatcher 

Occurs in east from Ivohibe north to Tsaratanana. Only a 
few records but reported as not uncommon. Dense humid 
forest. 170-1800 m. 

Terpsiphone mutata (R) Madagascar Paradise 

Flycatcher 
Widespread and common in all areas and variety of 
woodland habitat (included exploited forest). 0-2000 m. 
Also on Comoros, Seychelles, Mascarenes and Reunion. 

Family NECTARINIIDAE 
Nectarinia notata (R) Madagascar Green Sunbird 

Fairly common east and west forest and brush, including 
degraded woodland and central plateau. Less common in 
south-west. Prefers coastal plain to forested mountain 
slopes but found up to 1800 m. Less common than N. 
souimanga . Also Comoros. 

Nectarinia souimanga (R) Souimanga Sunbird 
Common in east and west in forest and brush, including 
degraded woodland. High altitudes bushes (2300 m in 
Tsaratanana massif) to subdesert undergrowth. Also on 
Aldabra, lies Glorieuses. 

Family ZOSTEROPIDAE 
Zosterops maderaspatana Madagascar White-eye 

Widespread and abundant in wooded areas throughout 
including exploited/degraded forest and afforestation areas. 



Madagascar Mannikin 



Family ESTRILDIDAE 
Lonchura nana (R) 
(= Lepidopygia nana ) 

Widespread and common, most records from east, north of 
Sahavandronina (Fianarantsoa) to Tsaratanana. Also in 
west as far south as Toliara. 0-2000 m. Primary forest, 
exploited, cultivated areas. Also on Mayotte. 

Family PLOCEIDAE 
Ploceus nelicourvi Nelicourvi Weaver 

East from Manombo north to north-west (Sambirano) and 
north-east; evergreen forest: unexploited, partially exploited 
and exploited. Fairly common. 0-1800 m. 

Foudia sakalava Sakalava Weaver 

Three discrete populations; localised, often absent from 
apparently suitable areas. South-west, north-centre 

(around Mahajanga), north-east (Antseranana): forest and 
wooded savanna, mangroves. Locally very common. 



■174- 






Foudia madagascariensis (R) Madagascar Fody 
West, east, subdesert. Common. 0-2000 m Especially in 
open brushy ground, remnant woodlands, clearings, 5 
afforestation areas. On government list of harmful species 
because of rice damage. Hybridizes with F. omissa . Also on 
Reunion and Mauritius, Rodrigues. 

Foudia omissa Forest Fody 

Restricted to humid east as far as Manombo and north 
(Antseranana); hybridizes with F. madagascariensis . 
Extends to central plateau (Manjakatompo); forest, 
sometimes degraded, rice fields. 0-1800 m. 



Madagascar Starling 



Family STURNIDAE 
Saroglossa aurata 
(= Hartlaubius auratus ) 
East, north, west as fas as Tsiandra; common, occurring in 
forest, secondary woodland, wooded savanna, wood edges, 
near cultivated areas. 0-1800 m. More common on coastal 
plain than inland. 

Family DICRURIDAE 

Dicrurus forficatus (R) Crested Drongo 

Widespread throughout wooded areas. Rather common even 
in open country, uses degraded woodland. Rare above 
1000 m, absent central plateau. Also Comoros. 



II. NON-ENDEMIC BIRD SPECIES 

(B) indicates breeding species 
(I) indicates introduced species 

Podicipedidae 

Tachvbaptus ruficollis (B) Little Grebe 
Common throughout and expanding. 



Diomedeidae 
Diomedea melanophris 
Very rare accidental to South 



Black-browed Albatross 



Procellariidae 

Macronectes giganteus Giant Petrel 

Accidental to east, south-east, south. 

Pachyptila vittata Broad-billed Prion 

Very rare accidental to east, north-east. 

Puffinua pacificus (B) Wedge-tailed Shearwater 

Rare breeder (on islands near Morombe); visitor to 
north-east. 

Hydrobatidae 

Oceanites oceanicus Wilson's Petrel 

Visitor to south, south-east and east; common Apr-July. 



Fregatta tropica 



Indian Ocean Black-bellied 
Petrel 
Accidental to south, south-east and east. 



Phaethontidae 
Phaethon aethereus 
Very rare accidental. 



Red-billed Tropicbird 



Phaethon lepturus (B) White-tailed Tropicbird 

Rare in south, south-east and west. Breeds Antseranana. 



Phaethon rubricauda 
Fairly regular accidental. 



Red-tailed Tropicbird 



Sulidae 

Sula sula Red -footed Booby 

Rare visitor; breeds lies Glorieuses, Europa, Tromelin. 

Sula leucogaster Brown Booby 

Rare visitor; breeds lies Glorieuses. 

Phalacrocoracidae 

Phalacrocorax africanus (B) Long-tailed Cormorant 
Rather common throughout; 0-1500 m. 



Appendix 2: faunal lists 

Anhingidae 
Anhinga rufa (B) African Darter 

Rather common throughout; 0-1400 m. 

Pelecanidae 
Pelecanus rufescens (B)? Pink-backed Pelican 

Occasional in west; may breed. 

Fregatidae 
Fregata minor Great Frigatebird 

Visitor to east and west coasts. 

Fregata ariel Lesser Frigatebird 

Very rare, accidental north, north-west and east. 

Ardeidae 
Ixobrvchus minutua (B) Little Bittern 

Occurs throughout but rather rare. 0-1000 m. 

Nvcticorax nvcticorax (B) Night Heron 

Rather common, throughout but localised. 0-1500 m. 

Butorides striatua (B) Green-backed Heron 

Rather common throughout. 0-750 m. Less common in 

west, becoming rarer in south. 

Ardeola ralloides (B) Squacco Heron 

Rather common throughout, particularly in west. 0-1800 m. 

Bubulcus ibia (B) Cattle Egret 

Common, found throughout. 0-1800 m. 

Egretta ardesiaca (B) Black Egret 

Rare and localised, north and centre. 

Egretta gularis (B) Western Reef Heron 

Widespread other than in dense forest or extensive savanna. 

Egretta alba (B) Great White Egret 

Rather rare though widespread breeder. 0-800 m. 

Ardea cinerea (B) Grey Heron 

Rather common throughout, except for east. 0-1400 m. 



Ardea melanocephala 
Rare straggler to West. 



Black-headed Heron 



Ardea purpurea (B) Purple Heron 

Common breeder, 0-1800 m. Normally away from coast. 

Ardea goliath Goliath Heron 

Very rare straggler to west. 

Scopidae 
Scopus umbretta (B) Hamerkop 

Common breeder throughout, 0-1800 m. Numbers declining 
for no obvious reason. 

Ciconiidae 

Anastomus lamelligerus (B) Openbill 
Rather common throughout except in east. 0-1200 m. 

Mvcteria ibis (B) Yellow-billed Stork 

Mainly west and south-west, also centre (especially Lake 

Alaotra). 

Threskiornithidae 
Plegadis falcinellus (B) Glossy Ibis 

Common throughout but numbers declining. 0-1500 m. 

Threskiornis aethiopica (B) Sacred Ibis 
Rather rare, rivers south-west, west, north-west. 

Platalea alba (B) African Spoonbill 

Rare breeder in west and north. 

Phoenicopteridae 
Phoenicopterus ruber (B) Greater Flamingo (B) 

Localised in west. Breed8 in L. Tsimanampetsotsa and L. 
Ihotry. 



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An environmental profile of Madagascar 



Phoeniconaias minor (B) Lesser Flamingo (B) 

Rather common south-west and west, breeds L. Ihotry. 

Anatidae 
Dendrocygna bicolor (B) Fulvous Tree Duck 

Rather common but localized; declining through hunting. 

Dendrocygna viduata (B) White-faced Tree Duck 

Common throughout, 0-1500 m; commonest duck on 
Madagascar. 

Sarkidiornis melanota (B) Knob-billed Goose 

Rather common especially in west. 0-1200 m. Declining 

throughout. 

Nettapus auritus (B) Dwarf Goose 

Rather common throughout. 

Anas erythrorhynchos (B) Red-billed Teal 
Common throughout, 0-1500 m. 

Anas hottentota (B) Hottentot Teal 

Rather common througout, 0-1800 m. 

Thalassornis leuconotus (B) White-backed Duck 

Rather rare though found throughout (except at altitude), 

0-1000 m. Lakes, marshes. 

Accipitridae 

Milvus migrans (B) Yellow-billed Kite 

Common throughout especially centre and west, less so east 
and south-west. 0-2000 m. 

Macheiramphu8 alcinus (B) Bat-eating Hawk 
Throughout forests but rather rare. Wooded savanna, dense 
forest with cliffs. 



Falconidae 
Falco eleonorae 



Eleanora's Falcon 



Rostratulidae 
Rostratula benghalensis (B) Painted Snipe 
Rather common breeder, though rare in South-west. 
0-1200 m. 

Recurvirostridae 

Himantopus himanotpus (B) Black-winged Stilt 
Rather common throughout, 0-750 m. No breeding colony 
yet located. 



Recurvirostra avosetta 
Accidental visitor. 



Common Avocet 



Dromadidae 
Dromas ardeola (B)? Crab Plover 

Rather common west, north-west and north coasts, found 
throughout year, probably breeds. 



Charadriidae 
Charadrius hiaticula 
Accidental migrant, Oct-Mar. 



Common Ringed Plover 



Charadrius pecuarius (B) Kittlitz's Plover 

Common breeder, west and south-west, less so in east. 

0-950 m. 

Charadrius marginatus (B) White-fronted Plover 
Rather common breeder. 

Charadrius tricollaris (B) Three-banded Plover 

Rather common throughout, 0-1800 m. 



Uncommon, winter visitor. Records from wide area. 



Charadrius mongolus 
Rare accidental. 

Charadrius leschenaultii 
Fairly common migrant. 

Pluvialis dominica 
Accidental visitor. 



Lesser Sand Plover 



Greater Sand Plover 



Lesser Golden-Plover 



Falco concolor 



Sooty Falcon 



Rather rare migratory visitor, (December-April) mostly in 
west and south-west. 

Falco peregrinus (B) Peregrine 

Resident, rare but throughout. 0-1000 m. Small endemic 

subspecies. 



Phasianidae 
Coturnix coturnix Common Quail 

Local, north, north-west, centre, east. 0-1500 m. 
from Africa, no definite evidence of breeding. 



Migrates 



Coturnix delegorguei (B) Harlequin Quail 

Breeds; common in north savanna, west and south-west. 

Open grasslands. 

Numididae 

Numida meleagris (I) Helmeted Guineafowl 

Common in west and south-west; somewhat less so in east. 
Declining through hunting. 

Rallidae 

Porzana pusilla (B) Baillons Crake 

Breeds, elusive and uncommon though found throughout in 
humid areas 0-1800 m. 

Porphvrio porphvrio (B) Purple Swamphen 

Widespread in wetlands and marshes. 

Porphvrio alleni (B) Allen's Swamphen 

Fairly common breeder in larger marshes, 0-750 m. 

Gallinula chloropus (B) Moorhen 

Common throughout, 0-1800 m. 

Fulica cristata (B) Red-knobbed Coot 

Rather common throughout. 0-1800 m. 



Pluvialis squatarola Grey Plover 

Fairly common migrant, Sept-Apr. 

Scolopacidae 
Calidris alba Sanderling 

Fairly common migrant, Sept-Apr. 



Calidria alpina Dunlin 

Rare visitor to coasts. 

Calidris minuta Little Stint 

Vagrant. 

Calidris ferruginea 

Fairly common migrant Oct-Mar. 

Philomachus pugnax Ruff 

Rare migrant. 



Curlew Sandpiper 



Limosa limosa 



Black-tailed Godwit 



Rare visitor, recorded north of Antananarivo. 



Limosa lapponica 
Rare migrant Nov-Mar. 



Bar-tailed Godwit 



Numenius phaeopus Whimbrel 

Common migrant Sept -April. 

Numenius arquata Eurasian Curlew 

Uncommon migrant Nov-April, has been seen throughout 

year. 

Tringa stagnatilis Marsh Sandpiper 

Uncommon migrant to freshwater. 

Tringa nebularia Common Greenshank 

Rare migrant, usually Nov-March, has been recorded 

June-Aug. 



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Appendix 2: faunal lists 



Tringa ochropus 
Vagrant. 



Green Sandpiper 



T ringa glareola Wood Sandpiper 

Rare migrant in West, Oct-Mar. 

Xenus cinereus Terek Sandpiper 

Fairly common migrant Oct-April. 

Actitis hypoleucos Common Sandpiper 

Common migrant Aug-Mar. 

Arenaria interpes Ruddy Turnstone 

Fairly common migrant Sept -May, some remain through 

year. 



Arctic Skua 
Great Skua 

Grey-headed Gull 

Silver Gull 

Southern Black-backed Gull 



Stercorariidae 
Stercorarius parasiticus 
Rare, west coast. 

Stercorarius skua 
No information. 

Laridae 

Larus cirrocephalus (B) 
Rare and local. 

Larus novaehollandiae 
No information. 

Larus dominicanus 

Rather common South-west coast. 

Gelochelidon nilotica Gull-billed Tern 

No information. 

Sterna caspia (B) Caspian Tern 

Widespread though rare and localised. 

Sterna bergii (B) Swift Tern 

Rather common on all coasts except east. Breeds on coastal 

islands. 

Sterna bengalensis (B)? ?Lesser Crested Tern 

Commonest tern on Madagascar. Probably breeds. 

Sterna dougallii (B) Roseate Tern 

Erratic visitor to north, north-east and south-west, often in 

considerable numbers. Breeds Toamasina, Toliara. 

Sterna hirundo Common Tern 

Visitor Dec-Jan. 

Sterna anaethetus (B) Bridled Tern 

Rare accidental. 

Sterna fuscata (B) Sooty Tern 

Breeds Toamasina, Toliara and lies Glorieuses. 

Sterna saundersi Saunder's Tern 

Rather common visitor, Nov-Mar. 

Chlidonias hvbrida (B) Whiskered Tern 

Rather common but localised, breeds Oct-Nov. 

Chlidonias leucopterus White-winged Black Tern 

Migrant visitor to lakes, Dec-Jan. 



Columbidae 
Oena capensis (B) Long-tailed Dove 

Common breeder throughout except East, fluctuating 
population and distribution. 0-1500 m. 

Cuculidae 
Pachycoccvx audeberti (B) Thick-billed Cuckoo 
Known only from 5 specimens all from East forest Sihanaka 
- Bay d' Antongil in most densely forested areas. Always 
uncommon, last collected June 1922. African counterpart 
can be easily overlooked, so it may still survive. 

Tytonidae 
Tyto alba (B) Barn Owl 

Rather common, east, north-east, also in west, subdesert. 
0-1800 m 

Asio capensis (B) Marsh Owl 

East, west into edge of subdesert. 0-1800 m. Quite common 

but local. 

Apodidae 
Apus melba (B) Alpine Swift 

Throughout, wide ranging and reasonably common, 
commonest in north-east. 0-1300 m. 



Apus barbatus (B) 
Breeds throughout. 



African Black Swift 



Cvpsiurus parvus (B) African Palm Swift 

Rather common, except high plateau and heavy forest. 

0-1100 m. 

Meropidae 
Merops superciliosus (B) Bee-eater 

Common throughout except high plateau and dense forest. 
Breeds. 

Upopidae 
Upupa epops (B) Hoopoe 

Distinctive endemic subspecies f U. e. marginata l: widespread 
except dense forest. 



Hirundinidae 
Riparia riparia 
Winter vagrant. 



Sand Martin 



Riparia paludicola (B) African Sand Martin 

Rather common breeder in east and High Plateau, especially 

at high altitude (500-2400 m). 

Hirundo rustica Swallow 

Irregular west coast winter visitor. 

Muscicapidae 
Saxicola torquata (B) Stonechat 

Common throughout, especially at altitude. 0-2400 m. 
Breeds. 

Sturnidae 

Acridotheres tristis (I) Indian Mynah 

Introduced, common and increasing. Competes with Upupa 
epops . Avoids altitude and forest; its expansion is an 
indication of deforestation. 



Anous tenuirostris 
Rather common on coasts. 



Lesser Noddy 



Corvidae 
Corvus albus (B) 
Common throughout. 



Pied Crow 



Anous stolidus (B) Brown Noddy 

Rather common on coasts, breeds south of Toamasina and 

lies Glorieuses. 

Gygis alba White Tern 

Found occasionally on coasts. 



•177- 



An environmental profile of Madagascar 

MADAGASCAR MAMMALS 

All species are endemic unless otherwise indicated. Lemurs are 
treated in detail in Appendix 3. 

INSECTIVORA 

Family TENRECIDAE 
Echinops telfairi 

Apparently widespread and abundant in the drier, western 
and southern parts of the island. 

Hemicentetes semispinosus 

Apparently relatively widespread in the eastern forested 
regions; apparently occurs at lower altitude than the 
following species. 

Hemicentetes nigriceps 

Eastern forests, apparently at higher altitude than the 

preceding. 

Setifer setosus 

Apparently widespread and abundant over much of the 

island. 

(The two specimens from the eastern rain forest, named as 
' Dasogale fontoynonti ' are now known to be aberrant Setifer 
setosus (M. Nicoll in litt. , 28.10.86)) 

Tenrec ecaudatus 

Widely distributed on the island; catholic habitat 
preferences; introduced to the Comoros, Seychelles, Reunion 
and Mauritius. 

Geogale aurita 

Recorded from the north-east (Fenoarivo Atsinanana) and 

south-west (Toliara and Morondava); Common at Beza 

Mahafaly. 

Limnogale mergulus 

Recorded from freshwater streams in eastern Madagascar, 
including Vohitra, Andranotobaka, Amborompotsy and 
Antsampandrono. Rare. 

Microgale brevicaudata 

Known from Mananara (Mahanara) on the north-east coast, 

75 km south of Vohemar; possibly a very localized species. 

Microgale cowani 

Recorded from the Ankafina forest in the eastern Betsileo 

region and at Andasibe (Analamazaotra). 

Microgale crassipes 

Known only from the type from Antananarivo (possibly 
collected from the region of Andrangoloaka, 70 km east of 
the capital). 

Microgale decarvi 

Known from near the cave of Andrahomana near Taolanaro. 

Microgale drouhardi 

Collected from the Antseranana region of northern 

Madagascar and the Andringitra area in the south-east. 

Microgale longicaudata 

Recorded from Ankafina, in eastern Betsileo. 

Microgale longirostris 

Known from Ampitambe in the south-east. 

Microgale majori 

Known from the Ankafina forest in eastern Betsileo. 

Microgale (Paramicrogale) occidentalis 
Maintirano, western Madagascar 

Microgale parvula 

Known from the type specimen collected on the Montagne 

d'Ambre. 



Microgale principula 

Recorded from Midongy-du-Sud, south-eastern Madagascar. 

Microgale proxilicaudata 

Antseranana, northern Madagascar (may be a subspecies of 

M. longicaudata ). 

Microgale pusilla 

Described from Ikongo forest near Vinanitelo; apparently not 
rare in museum collections, though reportedly only known 
from the type locality. 



near Andevorante 



Microgale sorella 

Recorded from Beforona forest, 

northern Madagascar. 

Microgale taiva 

Collected in the south-east at Ambohimitombo forest in the 
Tanala region, north-eastern Betsileo; also known from 
Perinet and Vondrozo. 

Microgale thomasi 

Collected from Ampitambe forest, northeastern Betsileo, 

Ivohimanitra forest (Ambohimanga) in southeastern 

Madagascar, Perinet, Vondrozo and the forest of 

Antrangolonka. 

Microgale melanorrhachis 

Recorded from Perinet and Ivohibe in the east. 

Microgale (Nesogale) dobsoni 

Collected in Nandesen forest, central Betsileo, also 
Manjakatompo (Ankaratra massif), Andringitra, 

Ambohimitambo, Ampitambe and Vinanintelo. 

Microgale fNesogale) talazaci 

Collected in forest of Ikongo near Vinanitelo (south of 
Fianarantsoa) and from Perinet, Maroantsetra, Analapa, 
Vondrozo and Andapa. 

Microgale (Leptogale) gracilis 

Reputedly rare, though possibly widespread in central 
eastern forests. Specimens from the forest of 

Ambohimitambo (eastern Betsileo), Ankeramadinika, 
Perinet and Andringitra. 

Orvzorictes hova 

Recorded from Ankaye and Antsianaka in central 

Madagascar; also Perinet and the Maroansetra region. 

Considerable confusion exists between this species and the 

following. 

Orvzorictes talpoides 

Recorded from the north-west (the Marovoay plain near I 

Mahajanga) and from the east (Maroantsetra and Perinet); I 

believed not to be rare, and to live in proximity to human I 

habitations. 

Orvzorictes (Nesorvctes) tetradactylus 

Recorded from Vinanitelo south-east of Fianarantsoa, the I 

high plateaux region, Antsirabe and Andringitra. 

Suncus etruscus f=madagascariensis1 

NON-ENDEMIC. North African coast and Ethiopia; I 
possibly also West Africa. Madagascan form (also on 1 
Comoros) may be separate species. 



Suncus murinus 






NON-ENDEMIC. North-east coast of Africa from Egypt to I 
Tanzania, also Zanzibar and Comoros. Also widespread in T 
Asia, where probably introduced. 

CHIROPTERA 

Family PTEROPODIDAE 

Eidolon helvum 

NON-ENDEMIC. Widespread in the Afrotropical region. 



■178- 



Appendix 2: faunal lists 



Pteropus rufus 

Madagascar; abundant in coastal regions, though otherwise 

localised (M. Nicoll, in litt. , 28.10.86). 

Rousettus madagascariensis 

Madagascar, apparently from the east; has been included in 
the widespread R. lanosus though its status as a separate 
species has been affirmed. 

Family EMBALLONURIDAE 
Emballonura atrata 

Eastern and central Madagascar. The sole member of the 
genus outside its range in south-east Asia, Malaysia and the 
South Pacific. 

Taphazous mauritianus 

NON-ENDEMIC. Widespread in the Afrotropical region. 

Family NYCTERIDAE 
Nycteris madagascariensis 

Madagascar. This may be the Madagascan representative of 
the widespread Nycteris thebaica . 

Family HIPPOSIDERIDAE 
Hipposideros commersoni 
NON-ENDEMIC. Widespread in the Afrotropical region. 

Triaenops furculus 

North and western Madagascar, also Aldabra. 

Triaenops humbloti 

Eastern Madagascar; may be only a colour variant of T. 

rufus . 

Triaenops persicus 

NON-ENDEMIC. Afrotropical region and Iran. 

Triaenops rufus 
Eastern Madagascar. 

Family MYZOPODIDAE 
Myzopoda aurita 

An endemic species in a monotypic family known from few 
specimens, mostly collected before 1900 and apparently from 
widely scattered localities mostly in the east of Madagascar 
(from Taolanaro north to Maroantsetra), though there is a 
record from Mahajanga in the west. 

Family VESPERTILIONIDAE 
Eptesicus capensis 
NON-ENDEMIC, widespread in the Afrotropical region. 

Myotis goudoti 

Madagascar, also Anjouan in the Comoros. 

Pipistrellus nanus 

NON-ENDEMIC, widespread in the Afrotropical region. 

Scotophilus nigrita 

NON-ENDEMIC, widespread in the Afrotropical region. 

Scotophilus robustus 

Madagascar. Considered by Hayman and Hill to be perhaps 
the Madagascan representative of the widespread S. nigrita , 
though its validity as a separate species has been reasserted. 

Scotophilus dinganii 

NON-ENDEMIC, widespread in the Afrotropical region; has 

been included in S. nigrita . 

Scotophilus barbonicus 

Madagacar and Reunion. Has been stated by Hayman and 
Hill to perhaps belong to the widespread S. leucogaste r, 
though its validity as a species has been reasserted. 

Miniopterus minor 

NON-ENDEMIC, widespread in the Afrotropical region. 



Miniopterus schreibersi 

NON-ENDEMIC; very wide distribution from southern 

Europe through Africa and Asia to Australia. 

Family MOLOSSIDAE 
Otomops martiensseni 

NON-ENDEMIC, occurs in eastern Africa from Djibouti to 
Angola and South Africa. 

Mormopterus acetabulosus 

NON-ENDEMIC, also known from Ethiopia, Reunion, 

Mauritius and South Africa. 

Mops condvlurus 

NON-ENDEMIC, widespread in the Afrotropical region. 

Mops midas 

NON-ENDEMIC, widespread in the Afrotropical region. 

Tadarida fulminans 

NON-ENDEMIC, known from east Africa, as far west as 

eastern Zaire. 

Mormopterus iugularis 

Madgascar. Genus has been included in Tadarida . 

Chaerophon pumila 

NON-ENDEMIC, widespread in the Afrotropical region. 

PRIMATES 

Family CHEIROGALEIDAE 
Allocebus trichotis 

Known only from four specimens collected from the eastern 
forests, most recently in 1965. 

Cheirogaleus major 

Reportedly fairly abundant in eastern rainforests. 

Cheirogaleus medius 

Still widespread in the dry, deciduous western and southern 

forests from the Bay of Narinda to Taolanaro. 

Microcebus fMirza) coquereli 

Restricted range in the forests of the west principally 
between the Onilahy and Fierenana rivers, and also on the 
Ampasindava Peninsula and the adjoining region. 

Microcebus murinus 

The most abundant lemur, along with M. rufus , occurring 
throughout the forested areas of the west, south and 
south-west from Taolanaro to the Sambirano region; also on 
Nosy Be. 

Microcebus rufus 

Occurs throughout the forests of the eastern region, from 
Fort-Dauphin to the Montagne d'Ambre. Has been 
considered a subspecies of M. murinus . 

Phaner furcifer 

A specialized gum-feeder, occurring principally in the west, 

though with email populations in the north, east and south. 

Family INDRIIDAE 
Avahi laniger 
Occurs in the eastern and north-western forests. 

Indri indri 

Restricted to coastal and montane rainforest from sea level 
to around 1800 m in north-eastern Madagascar, from 
Antongil Bay in the north to the Masora River in the south. 
The largest lemur. 

Propithecus diadema 

Found in forests of the north-east and east. 

Propithecus verreauxi 

Has a wide distribution in the north-west, west, south-west 

and south. 



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An environmental profile of Madagascar 



Family DAUBENTONIIDAE 
Daubentonia madagascariensis 

Believed to be more widespread than previously thought in 
the eastern forests; introduced to Nosy Mangabe where still 
survives. 

Family LEMURIDAE 
Hapalemur simus 

Apparently known only from humid forest in the 
Fianarantsoa region, where recently recorded at Ranomafana. 



Hapalemur griseus 

Found in eastern and north-western 

occurring over a wide altitudinal range. 



forested regions, 



Lemur catta 

Occurs in the dry forests of the south and south-west where 

it is considered still relatively abundant. 

Lemur coronatus 

Found in the extreme north of the island as far south in the 
west as the Ankarana limestone karat and in the east to the 
Fanambana River. 

Lemur fulvus 

Widespread in forested regions of the island other than in 

the extreme south; has been considered conspecific with L. 

macaco but populations of the two are now known to exist in 

sympatry in the north and specific distinction is generally 

upheld. 

Lemur macaco 

Found in humid forests of the north-west (including the 
Tsataratanana Massif) and the coastal islands of Nosy Be 
and Nosy Komba. 

Lemur mongoz 

Occurs in north-western Madagascar and on the Comorian 
islands of Moili and Ndzouani, to which it has almost 
certainly been introduced. 

Lemur rubriventer 

Appears to occur throughout the forested interior of the east 
from the Tsaratanana Massif in the north to the Andringitra 
massif in the south. 

Lepilemur mustelinus 

Found in the northern part of the eastern forests between 

Toamasina and Antalaha. 

Lepilemur microdon 

Occurs in the southern part of the eastern forests from 

Perinet to Taolanaro. 

Lepilemur leucopus 

Found in the dry southern forests from Taolanaro westwards 

possibly as far as the Onilahy River. 

Lepilemur septentrionalis 

Occurs in the extreme north, north of Ambilob£ and to the 

south and east of the Montagne d'Ambre. 

Lepilemur ruficaudatus 

Occurs in the western forests, though limits of range are 
ill-defined; to the south it occurs at least as far as the 
Onilahy River and to the north the boundary with L. 
edwardsi appears to be the Tsiribihina River. 

Lepilemur dorsalis 

Occurs in the Lokobe forest on Nosy Bi Island and the 

forests on the north-western coast of Madagascar facing this. 

Lepilemur edwardsi 

Confined to western Madagascar, from the Bay of 
Mahajamba south at least as far as Antsalova and possibly 
to the Tsiribihina River. 

Varecia variegata 

Occurs in the humid eastern rainforests. 



CARNIVORA 

Family VIVERRIDAE 
Fossa fossana 

Apparently widely distributed though at low density in 
wooded or forested areas in the east. 

Eupleres goudotii 

Rarely recorded and little known, though reported from the 
eastern coastal region and the north of the island; known to 
occur at Perinet. 

Galidia elegans 

Apparently occurs in wooded areas in the east and west. 

Mungotictis lineatus 

Inhabits deciduous woodlands on sand in the west and 
south-west of Madagascar; it is apparently relatively 
common, even in somewhat degraded areas. 

Galidictis striata 

Occurs in the eastern rainforests where it is apparently very 

little known, though according to reports not uncommon. 

Salanoia concolor 

Known from the eastern rainforests, where it is thought to be 

rather rare. 

Cryptoprocta ferox 

Widespread in Madagascar, though apparenly very rare on 

the central plateau. 

RODENTIA 

Family CRICETIDAE 
Hypogeomvs antimena 

Known only from the sandy coastal area in the region of 
Morondava. Burrows and is largely nocturnal. 






Macrotarsomys bastardi 

Found in dry regions in most of the western part of 

Madagascar; seems to be limited by the 20 * isotherm. 

Macrotarsomys ingens 

Known from the forest of Ankarafantsika in the north-west 
of Madagascar; also represented by fossil remains in the 
south-west of the island. 

Nesomvs rufus 

Eastern rainforests; one specimen was caught in 1928 at 

Maintirano on the western coast. Diurnal. 

Eliurus mino r 

Eastern rain forests. Originally described from Ampitambe 
forest in northeastern Betsileo; recently collected at Perinet 
and in the Maroansetra region. The Eliurus complex is 
highly variable and probably contains more than two species 
(M. Nicoll in litt. , 28.10.86). 

Eliurus mvoxinus 

Apparently widespread; believed likely to inhabit most of the 
residual forests in the west and centre of the island as well as 
the eastern rainforest. 

Gymnuromys roberti 

Eastern rain forests; originally described from Ampitambe 
forest in northeastern Betsileo. Seldom caught where Rattus 
rattus is locally abundant. 

Brachytarsomys albicauda 

Eastern forest species; strictly arboreal. 

Brachyuromys betailoensis 

Recorded from the south-eastern part of the Betsileo region, 
where it is known from some dozen specimens, and from the 
Andringitra massif. 

Brachyuromys ramirohitra 

Recorded from Ampitambe forest in the Betsileo region and 

from the Andringitra massif. 



180- 






Appendix 2: fauna! lists 



MADAGASCAR REPTILES 

NB. Provisional IUCN categories are cited (see Part V.3); 
those with an asterisk are quoted from a published IUCN Red 
Data Book - these species are treated in detail in Appendix 3. 
Cardinal points are abbreviated N, S, E, W and C (central). 
Madagascan names, where known, are quoted in single 
inverted commas. For references see Part V.3. 

TESTUDINES 

Family TESTUDINAE 
Geochelone radiata Shaw, 1802 V* 

ENDEMIC. Restricted to Didierea forest occurring in a 
narrow arc across southern Madagascar; has been recorded 
from near Amboasary in the south-east to near Morombe in 
the south-west. 'Sokake'. 

Geochelone vniphora Vaillant, 1885 E* 

ENDEMIC. Restricted to three forest 'islands' in the 
vicinity of Baly Bay, including Cape Sada, in northwest 
Madagascar. 'Angonoka'. 

Kinvxis belliana Gray, 1831 

Introduced; acclimatised in the northwest, also recorded near 

Amboasary in the southeast. 

Pyxis arachnoides Bell. 1827 I* 

ENDEMIC. Restricted to coastal areas (10- 20km inland) in 
the south and south-west, from Morombe in the south-west 
to Amboasary in the south-east. 

Pyxis planicauda (Grandidier, 1867) I* (prob E) 
ENDEMIC. Apparently restricted to the Andranomena 
forest, an area of approximately 100 sq km situated 20km 
northeast of Morondava on the central-west coast. 
'Kapidolo'. 



Pelusios castanoides Hewitt, 1931 

Almost throughout the western half of Madagascar and 
along coastal areas in the southeast and east. Eaten locally. 
Widespread in eastern Africa. 

Pelusios subniger (Lacepede, 1788) 

Occurs along the eastern littoral; possibly an ancient 
introduction. Eaten locally. Widespread in south-central 
and eastern Africa. 

CROCODYLIA 

Family CROCODILIDAE 
Crocodvlus niloticus (Laurenti 1768) V* 

Reported common in most waters during the nineteenth 
century, now very diffuse and rare due to persecution and 
over-exploitation. Widespread in Africa. 'Voay' or 
'Mamba'. 

SAURIA 



Family GEKKONIDAE 
Ailuronvx trachygaster (Dumeril, 
ENDEMIC. Locality unknown. 



1851) 



Ebenavia inunguis Boettger, 1878 

NE, E, SE, SW, St. Marie, Nosy Be, Nosy Mangabe (etc); 
also on Comoros and Mauritius. In Reserve du Betampona 
(Toamasina region). On coastal rocks, under bark, in 
forests, to 800m. Common on St. Marie. 

Geckolepis anomala Mocquard, 1909 
ENDEMIC. S. 



Geckolepis maculata Peters, 1880 
NW, E, W, Nosy Be; also Comoros, 
tree trunks. 



Found in huts, cracks in 



Family CHELONIIDAE 
Caretta caretta (Linnaeus, 1758) V* 

Nesting occurs particularly in the southeast, around 
Taolanaro, and with some nesting along the west coast as far 
north as Morondava. Annual nesting numbers estimated at 
less than 300. Under pressure from exploitation. 

'Ampombo'. 

Chelonia mvdaa (Linnaeus, 1758) E* 

Small scale nesting occurs. Exploited, mainly for domestic 

consumption. 'Fanonjato'. 

Eretmochelys imbricata (Linnaeus, 1766) E* 
Good numbers still nest, mainly in the northern third of the 
island and in the southwest, but is heavily exploited and 
populations have declined to a remnant of former size. An 
estimated 2500 Hawksbills are taken annually, mainly by the 
Ve«o people of the southwest. 'Fano'. 

Lepidochelys olivacea (EschscholU, 1829) E* 

Nesting has been recorded in the northwest although other 

reports suggest presence of a feeding population only. 

Family DERMOCHELYIDAE 
Dermochelvs coriacea (Vandelli, 1761) E* 
Rare, accidental in Madagascar waters. 

Family PELOMEDUSIDAE 

Ervmnochelys madagascariensis (Grandidier, 1867) I* 
ENDEMIC. Present in freshwater habitats at low to 
moderate altitude in the west and northwest, from the 
Mangoky River in the southwest, northward to the 
Sambirano basin west of the Massif de Tsaratanana. 
Exploited for food by the riverine Sakalava people and 
others around Lake Kinkony, possibly affected by habitat 
loss. 'Rer«'. 



Pelomedusa subrufa (Lacipede, 1788) 
Present almost throughout Madagascar. 
Widespread in Africa. 'Kapika'. 



Geckolepis petiti Angel, 1942 

ENDEMIC. Andranovaho, Mahafaly Prov. Type locality 

only. 

Geckolepis polylepis Boettger, 1893 
ENDEMIC. NW, SW, W, St.Marie. 



Geckolepis typica Grandidier, 1867 
ENDEMIC. NE, SE, SW. 

Gehvra mutilata (Wiegmann, 1835) 

E; very widespread outside Madagascar. 

frequenting human habitation. 



Well known for 



Hemidactylus mabouia (Moreau de Jonnes, 1818) 

Common throughout; very widespread outside Madagascar. 

In huts, on coast, low mountains. 

Hemidactylus frenatus Dumeril et Bibron, 1837 

Common throughout; very widespread outside Madagascar. 



' Hemidactylus gardinieri ' Boulenger, 1909 

Listed as synonym of H. mercatorius Gray 1842 

Tierreich Gekkonidae checklist; widespread 

Madagascar. 



in Das 
outside 



Not eaten. 



Homopholis antongilensis Bohme & Meier, 1980 
ENDEMIC. Antongil Bay area, including Nosy Mangabe. 

Homopholis boivini (Dumeril, 1856) 

ENDEMIC. Extreme north, between Ambilobe and 
Antseranana (Diego Suaree). (Removed from synonymy of 
H. heterolepis , Bohme & Meier, 1980.). 

Homopholis sakalava (Grandidier, 1867) 

ENDEMIC. Arid SW and W, Ampanihy to Mahajanga. 
(Removed from synonymy of H. heterolepis (sensu Russell, 
1978), Bohme & Meier, 1980; includes in synonymy H. 
heterolepis Boulenger, 1896. 



■181- 



An environmental profile of Madagascar 



Lygodactvlus arnoulti Pasteur, 1964 

ENDEMIC. C - Montagne de l'lbity, 25km S of Antsirabe, 

2150m. Type locality only. Under stones. 

Lygodactvlus blanci Pasteur, 1967 
ENDEMIC. C - Mt. Ibity. 

Lygodactvlus cowani 

ENDEMIC. SW, C, S-C. Synonym: Microscalabotes bivittis 

(Peters, 1883). 

Lygodactvlus decaryi Angel, 1930 

ENDEMIC. SE - Massif de l'Angavo, Taolanaro Prov. 

Single specimen only, collected under bark, 400m. 

Lygodactvlus expectatus Pasteur & Blanc, 1967 
ENDEMIC. Ambilobe karst (Ankarana massif), 12km NNW 
of Ankarana also Antseranana area. Among masses of fallen 
debris at foot of karst cliffs, in dry vegetation, on rocks. 
Partly or totally non-arboreal. 

Lygodactvlus guibei Pasteur, 1964 

ENDEMIC. E - Pgrinet, between 

Toamasina-Antananarivo. Arboreal. In humid forest. 

Lygodactvlus heterurus Boettger, 1913 
ENDEMIC. Nosy Be. 

Lygodactvlus klemmeri Pasteur, 1964 
ENDEMIC. NW - Foret de l'Antsingy. 

Lygodactvlus madagascariensis (Boettger, 1881) 

ENDEMIC. NE, E, SE., inc. Mt. d'Ambre & Sambirano. 

Arboreal, in humid forest, to 1000m, also rocks, scrub. 



Phelsuma barbouri Loveridge, 1942 

ENDEMIC. E - Forest between Toamasina-Antananarivo, 
also Tsiafajavona (main summit of Ankaratra massif, SSW 
of Antananarivo). Arboreal, also on & among rocks on 
Tsiafajavona. Feeds on small invertebrates. 

Phelsuma bimaculata Kaudern, 1922 
ENDEMIC. E. 

Phelsuma dubia (Boettger, 1881) 

NW, SW, W, S-C, Nosy Be; also in Tanzania. 'Sasaka'. 



c 100km E of 



Lygodactvlus miops Giinther, 1891 
ENDEMIC. E, SE. East of Antalaha. 
forest. 



Arboreal. In humid 



Phelsuma flavigularis Mertens, 1962 
ENDEMIC. E - Perinet (950m) 

Antananarivo. Type locality only. 

Phelsuma guttata Kaudern, 1922 
ENDEMIC. NE, E, S. 



Phelsuma laticauda (Boettger, 1880) 

NW, SE, S, SW, W, Nosy Be; also Comoros and Seychelles. 

Phelsuma lineata Gray, 1831 

ENDEMIC. NW, NE, E, SE, C, S-C. Moderately common 
forest species, found in coastal areas & up to 1100 m, on 
Agave leaves around Toamasina. Maximum density along 
east coast. 'Antsiantsy'. 

Phelsuma madagascariensis Gray, 1831 

NW, N, NE, E, S, SW, W, St. Marie, Nosy Be; also 
Seychelles. Very common in Antseranana, common in east 
and north generally. Enters huts, along littoral in SE, 
frequent in NW forests. 

Phelsuma mutabilis (Grandidier, 1869) 

ENDEMIC. NW, SE, S, SW, W, C. Very common in forest 
along River Onilahy. Found under bark, also in arid scrub. 
One of smallest Phelsuma , only 3cm snout-vent. 'Tsatsake'. 



Lygodactvlus montanus Pasteur, 1964 
ENDEMIC. SE-C - summit of Mt. 
specimens only. 



Ivohibe. 



Three 



Lygodactvlus ornatus Pasteur, 1964 
ENDEMIC. NW - Mt. Mandritsara. 
only. 



Known by holotype 



Lygodactvlus rarus Pasteur & Blanc, 1973 
ENDEMIC. Eastern cliffs of Ambilobe karst (extreme NE of 
Ankarana massif) and Mangindramo, and edges of 
Tsaratanana forested massif 1350m. 



Phelsuma quadriocellata (Peters, 1883) 
ENDEMIC. S, C. 

Phelsuma serraticauda Mertens, 1963 

ENDEMIC. E - Ivoloina, 12km N of Toamasina. 

locality only. 



Type 



Phelsuma standingi Methuen et Hewitt, 1913 
ENDEMIC. SW: Maroamalona, forests along R. Onilahy. 

Phelsuma trilineatum Gray, 1842 
ENDEMIC. SW. Locality unknown. 



Lygodactvlus robustus Boettger, 1913 
ENDEMIC. S, SE, S-C. 



Phvllodactvlus androyensis Grandidier, 1867 
ENDEMIC. SE, SW, St. Marie. Sunny rocks on coast. 



Lygodactvlus spinulifer 

ENDEMIC. NE, E, SE. Forest species, to 1000m. 

Lygodactvlus tolampyae (Grandidier, 1872) 
ENDEMIC. W, NW. 

Lygodactvlus tuberifer Boettger, 1913 
ENDEMIC. SW, W. 

Lygodactvlus tuberosus Mertens, 1965 
ENDEMIC. Localities unknown. 

Lygodactvlus verticillatus Mocquard, 1895 

ENDEMIC. SE, S, SW. On calcareous littoral rocks, scrub, 

bushes. 

Millotisaurus mirabilis Pasteur, 1962 

ENDEMIC. Mt. Tsiafajavona, 2300-2500m. Single locality. 

Paragehyra petiti Angel, 1929 

ENDEMIC. Lavenombato in Toliara Prov., at foot of 

calcareous cliffs in Mahafaly area. Single locality? 



Phvllodactvlus barbouri Angel, 1936 
ENDEMIC. 'Madagascar', no precise locality. 

Phvllodactvlus bastardi (Mocquard, 1900) 
ENDEMIC. SE, S, SW, W. On sunny rocks on coast, arid 
scrub, subdesert, to 350m. After P. pictus is most frequently 
seen species of genus. 

Phvllodactvlus gracilis (Boulenger, 1896) 
ENDEMIC. SW. 

Phvllodactvlus homalorhinus Angel, 1936 

ENDEMIC. N - Ankaratra, Ambilobe Dist., Antseranana 

Prov. Type locality only. 

Phvllodactvlus oviceps Boettger, 1881 

ENDEMIC. Nosy Be, Sakatia (islet W of Nosy Be). 

Phvllodactvlus pictus (Peters, 1854) 

ENDEMIC. SE, S, SW, W. Most frequently seen species of 

genus. On calcareous rocks on coast, etc. 

Phvllodactvlus porphyreus (Daudin, 1803) 

Rarely reported, localities unknown, presence requires 

confirmation; occurs in southern Africa. 



182- 



Appendix 2: fauna! lists 



Phvllodactvlus stumpffi Boettger, 1878-79 
ENDEMIC. N, Nosy Be. 

Uroplatus fimbriatus (Schneider, 1797) 

ENDEMIC. NE, E, SE, C, S-C, W-C, Ste. Marie, Nosy 

Mangabe, Nosy Be. Forest species, eats insects, 300-1100m. 

'Taha-fisaka'. 

Uroplatus lineatus Dumeril et Bibron, 1836 
ENDEMIC. E - Toamasina region. Taha-fisaka'. 

Uroplatus giintheri Mocquard, 1908 

ENDEMIC. Locality unknown. Single specimen. 

Uroplatus alluaudi Mocquard, 1894 

ENDEMIC. N - Mt. d'Ambre. Two specimens only. 

Uroplatus phantasticus (Boulenger, 1888) 
ENDEMIC. E, SE, S-C. 

Uroplatus ebenaui (Boettger, 1879) 
ENDEMIC. N, E, Nosy Be. Inc. Mt. d'Ambre. 

Family IGUANIDAE 
Chalarodon madagascariensis Peters, 1854 
ENDEMIC. SE, S, SW, W. Especially in S, less frequent in 
W. Very common in Ambovombe area and elsewhere. 
Sunny areas, on sand, in clearings in scrub forest. 
'Dangalia', 'Dangara'. 

Opiums cuvieri (Gray, 1831) 

NW, W; also Comoros. Common throughout NW. In 
savannah, in very dry regions, bushes. Formerly known as 
Opiums sebae Dumeril et Bibron, 1837 - see Savage 1952. 
'Sitry', 'Androngo'. 

Oplurus cyclurus (Merrem, 1820) 

ENDEMIC. SE, S, SW. 'Sitry', 'Androngo', 'Androngohazo'. 

Oplurus fierinensis Grandidier, 1869 

ENDEMIC. SW - Mahafaly area. 'Sitry', 'Androngo'. 

Oplurus grandidieri Mocquard, 1900 

ENDEMIC. E - Vinanitelo forest & Massif de l'lkongo. 

'Sitry', 'Androngo'. 

Oplurus quadrimaculatus Dumeril, 1851 

ENDEMIC. SE, S, SW. In sunny areas, rock, scrub, coast 

to 1800m. 'Sitry', 'Androngo'. 



Oplurus saxicola Grandidier, 1869 

ENDEMIC. SE, S, SW, S-C. Habitat inc. 

'Sitry', 'Androngo'. 



arid scrub. 



Family CHAMAELEONTIDAE 

NB: categories in this family fide Brygoo (in litt., 20.5.83.). 

Brookesia bonsi Ramanantsoa, 1980 K 

ENDEMIC. NW - In reserve naturelle No. 8, Tsingy de 

Namoroka, sub-prefecture of Soalala. 

Brookesia decaryi Angel, 1938 R 

ENDEMIC. W - coast, forests on Ankarafantsika massif. 



Brookesia dentata Mocquard, 1900 
ENDEMIC. NW - Suberbieville, 
Apparently known by Type only. 



S of Maevatanana. 



Brookesia ebenaui (Boettger, 1880) R 
ENDEMIC. NW, N, NE, E, C, S-C, Nosy Be. 

Brookesia griveaudi Brygoo, Blanc & Domergue, 1974 K 
ENDEMIC. NE. - Marojejy. 

Brookesia karchei Brygoo, Blanc &. Domergue, 1970 K 
ENDEMIC. NE - Mt Marojejy (in Reserve Naturelle). 
700m, dense shady high forest of the east region, on the 
ground on dead leaves, in areas of permanent humidity. 



Brookesia lambertoni Brygoo & Domergue, 1970 
ENDEMIC. Fito, in Sihanaka country. Type locality only. 

Brookesia legendrei Ramanantsoa, 1980 K 
ENDEMIC. Nosy Be, reserve naturelle No. 6. 

Brookesia minima Boettger, 1893 K 
ENDEMIC. Nosy Be. 

Brookesia nasus Boulenger, 1887 R 
ENDEMIC. E, SE, S-C. 

Brookesia perarmata (Angel, 1933) K 
ENDEMIC. Antsingy region, Menabe Prov., 300m. 

Brookesia pevrieresi Brygoo & Domergue, 1975 K 
ENDEMIC. NE - Nosy Mangabe, Antongil Bay. 

Brookesia ramanantsoai Brygoo & Domergue, 1975 K 
ENDEMIC. C - Ambohiboataba forest, east of Mantasoa. 

Brookesia stumpffi Boettger, 1879 nt 

ENDEMIC. NW, N, E, SW, W, Nosy Be. In forests, under 

dry leaves, on old rotten tree trunks. 

Brookesia superciliaris (Kuhl, 1820) nt 
ENDEMIC. NW, NE, E, SE, E-C, S-C, Nosy Be. 



Brookesia therezieni Brygoo & Domergue, 1970 K 
ENDEMIC. E, Perinet, Moramanga sub-prefecture, 
locality only. 



Type 



Brookesia thieli Brygoo & Domergue, 1969 K 
ENDEMIC. E. Moramanga and Maroantsetra 

sub-prefectures, inc. Perinet. A forest species, 900- 1500m, 
on the ground or among lichens on bushes, diurnal. 

Brookesia tuberculata Mocquard, 1894 R 
ENDEMIC. N - Mt. d'Ambre. 

Brookesia vadoni Brygoo & Domergue, 1968 K 

ENDEMIC. NW. Valley of the Iaraka River, near Masoala. 

600- 1000m. On branches of bushes. 

Chamaeleo angeli Brygoo & Domergue, 1968 K 

ENDEMIC. NW. On RN 4, N of Tsaramandroso (foret de 

1' Ankarafantsika). 

Chamaeleo antimena Grandidier, 1872 I 

ENDEMIC. SW. Recently removed from synonymy of C. 

rhinoceratus (Brygoo & Domergue, 1968). 

Chamaeleo balteatus Dumeril & Bibron, 1851 R 
ENDEMIC. Type locality unknown. Five new specimens 
from forest region between Ifanadiana and Fort Carnot. 
Recently removed from synonymy of C. bifidus (Brygoo & 
Domergue, 1969). 

Chamaeleo belalandaensis Brygoo & Domergue, 1970 
ENDEMIC. SW. Belalanda, 4 km N of Toliara. Single 
specimen. 

Chamaeleo bifidus Brongniart, 1800 R 
ENDEMIC. E, S-C. 

Chamaeleo boettgeri Boulenger, 1888 R 
ENDEMIC. N, Nosy Be. 

Chamaeleo brevicornis Glinther, 1879 nt 
ENDEMIC. C,S-C. 

Chamaeleo campani Grandidier, 1872 I 
ENDEMIC. C, E-C. 

Chamaeleo capuroni Brygoo, Blanc & Domergue, 1972 K 
ENDEMIC. 

Chamaeleo cucullatus Gray, 1831 R 
ENDEMIC. E. 



183- 



An environmental profile of Madagascar 



Chamaeleo fallax Mocquard. 1900 I 
ENDEMIC. E, SE, C, S-C. 

Chamaeleo furcifer Vaillant et Grandidier, 1880 
ENDEMIC. E, C. Known by holotype only. 

Chamaeleo gallus Giinther, 1877 R 
ENDEMIC. E, S-C. 

Chamaeleo gastrotaenia Boulenger, 1888 I 
ENDEMIC. NE, NW, E, C, S-C. 

Chamaeleo globifer Giinther, 1879 R 
ENDEMIC. C.S-C. 

Chamaeleo guibei Hillenius, 1959 

ENDEMIC. NE Tsaratanana and C-E "Sihanaka". Known 

by types only. 

Chamaeleo labordi Grandidier, 1872 K 

ENDEMIC. SW. Locally common along the Ihotry. 
Recently removed from synonymy of C. rhinoceratus 
(Brygoo & Domergue, 1968). 

Chamaeleo lateralis Gray, 1831 nt 

ENDEMIC. NE, E, SE, S, SW, W, C, S-C. 'Tanata', 

'Tanala', 'Tanalahy', 'Sangorita'. 

Chamaeleo linotus Miiller, 1924 K 

ENDEMIC. NE - Ambatodradama, 1000m, Maroantsetra 

Prov. 

Chamaeleo malthe Giinther, 1879 R 
ENDEMIC. N, NE, E, SE, C, S-C. 

Chamaeleo minor (Giinther, 1879) R 
ENDEMIC. S-C. 

Chamaeleo monoceras Boettger, 1913 

ENDEMIC. NW - Betsako near Mahajanga. Taxonomic 

status uncertain. Known by holotype only. 

Chamaeleo nasutus Dumeril et Bibron, 1836 nt 
ENDEMIC. N, NE, E, SE, C, S-C, St. Marie. 

Chamaeleo oshaughnessvi Giinther, 1881 K 
ENDEMIC. NE, E, S-C. 

Chamaeleo oustaleti Mocquard, 1894 nt 

(ENDEMIC). NW, N, NE, SW, C; & introduced to Ngong 

Forest near Nairobi, Kenya. 'Tana', 'Sangorita'. 

Chamaeleo pardalis Cuvier, 1829 nt 

(inc. C. guentheri (Boulenger, 1888)). (ENDEMIC). NW, 

N, E, inc Mt. d'Ambre; & introduced on Reunion. 

Chamaeleo parsonii Cuvier, 1824 I 
ENDEMIC. E, C, S-C, St. Marie, Nosy Be. 

Chamaeleo petted (Brygoo & Domergue, 1966) 
ENDEMIC. N - Ankarana massif (cf Ramanantsoa, 1978). 

Chamaeleo pevrieresi Brygoo, Blanc & Domergue, 1974 K 
ENDEMIC. 

Chamaeleo rhinoceratus (Gray, 1843) K 

(inc. C. voelukowi (Boettger, 1893)). ENDEMIC. NW, SW, 

W. 

Chamaeleo tsaratananensis Brygoo & Domergue, 1968 K 
ENDEMIC. N. Tsaratanana massif. 

Chamaeleo tuzetae Brygoo, Bourgat & Domergue, 1972 
ENDEMIC. SW - Andrenalamivola, near Ambiky, canton of 
Befandriana S. Known by holotype only. 

Chamaeleo verrucosus Cuvier, 1829 nt 
ENDEMIC. NW, N, E, SE, S, SW, W. 



Chamaeleo willsi Giinther. 1890 R 
ENDEMIC. C, E-C. 

Family SCINCIDAE 

NB: categories in this family fide Brygoo in litt., 20.5.83. 
Cryptoblepharus boutonii (Desjardins, 1831) nt 
Widespread; also very widespread outside Madagascar. 
Formerly assigned to Ablepharus , see Fuhn 1961. 

Amphiglossus andranovahensis (Angel, 1933) 

ENDEMIC. SW. Known by holotype only. Removed from 

Scelotes : Brygoo, 1984. 

Amphiglossus ankodabensis Angel, 1930 

ENDEMIC. SE. Only two specimens known. Removed 

from Scelotes : Brygoo, 1984. 

Amphiglossus ardouini (Mocquard, 1897) K 

ENDEMIC. N - Antseranana and Ambilobe. Removed from 

Scelotes : Brygoo, 1982. 

Amphiglossus astrolabi Dumeril et Bibron, 1839 K 
ENDEMIC. NE, E, S-C. 

Amphiglossus decarvi (Angel, 1930) 

ENDEMIC. SE. On littoral rocks. Known by holotype 

only. Removed from Scelotes : Brygoo, 1985. 

Amphiglossus frontoparietalis (Boulenger, 1889) K 
ENDEMIC. NE, E, SE. Synonym (Brygoo 1980): Scelotes 
praeornatus Angel, 1938. Removed from Scelotes : Brygoo 
1981. 

Amphiglossus gastrostictus (O'Shaughnessy, 1879) K 
ENDEMIC. E. Removed from Scelotes : Brygoo, 1984. 

Amphiglossus igneocaudatus (Grandidier, 1867) nt 
ENDEMIC. S, Centre. Removed from Scelotes : Brygoo, 
1984. 

Amphiglossus intermedius (Boettger, 1913) K 

ENDEMIC. N, W, S. Removed from Scelotes : Brygoo, 1984. 

Amphiglossus macrocercus (Giinther, 1882) K 

ENDEMIC. C, E, SE. Removed from Scelotes : Brygoo. 1984. 

Amphiglossus macrolepis (Boulenger, 1888) 

ENDEMIC. SE - Taolanaro. Only two specimens known. 

Removed from Scelotes : Brygoo, 1981. 

Amphiglossus melanopleura (Giinther, 1877) K 
ENDEMIC. N, E. In forest. Removed from Scelotes : 
Brygoo, 1982. 

Amphiglossus melanurus (Giinther, 1877) nt 

ENDEMIC. Widespread, C, E, S. Removed from Scelotes : 

Brygoo, 1984. 'Matahotandro', 'Ankotofotsy'. 

Amphiglossus mouroundavae (Gradidier, 1872) K 
ENDEMIC. N, W. In humid areas, under stones, in tree 
trunks. Removed from Scelotes : Brygoo, 1984. 

Amphiglossus ornaticeps (Boulenger, 1896) R 
ENDEMIC. S, E. Removed from Scelotes : Brygoo, 1984. 

Amphiglossus poecilopus (Barbour & Loveridge, 1928) K 
ENDEMIC. Widespread, E, S, W. Removed from Scelotes : 
Brygoo, 1984. 

Amphiglossus polleni (Grandidier, 1869) K 

ENDEMIC. NW, E, SW, W, Nosy Be. Removed from 

Scelotes : Brygoo, 1982. 

Amphiglossus reticulatus (Kaudern, 1922) 

ENDEMIC. NW. Known by type only. Removed from 

Scelotes : Brygoo, 1980. 

Amphiglossus splendidus (Grandidier, 1872) R 

ENDEMIC. NW, NE, E, SE, C. Removed from Scelotes : 

Brygoo, 1982. 



184- 



Amphiglossus stumpffi (Boettger, 1882) E 

ENDEMIC. Nosy Be. Removed from Scelotes: Brygoo, 1982. 

Amphiglossus tsaratananensis Brygoo, 1981 

ENDEMIC. N - Tsaratanana Mtns. Known by types only. 

Removed from Scelotes : Brygoo, 1982. 



Amphiglossus waterloti (Angel, 1930) K 
ENDEMIC. N - Ambilombe, Antseranana area; 
Antsohihy area. 



also Bora, 



Androngo allaudi (Brygoo, 1981) 

ENDEMIC. N. Mt d'Ambre, Ankarana, Antseranana. 

Known by types only. Removed from Scelotes : Brygoo, 1982. 

Androngo crenni Mocquard, 1906 

ENDEMIC. E. Only two specimens known. Removed from 

Scelotes : Brygoo, 1982. 

Androngo elongatus Angel, 1933 K 

ENDEMIC. N - Ambilombe, Antseranana area, Nosy Be. 

Removed from Scelotes : Brygoo, 1982. 

Androngo trivittatus (Boulenger, 1896) R 

ENDEMIC. S. A species of the hot dry extreme south. 

Synonyms (Brygoo, 1979): Pygomeles trivittatus , S^ 

trilineatus Angel, 1949. Removed from Scelotes : Brygoo, 

1982. 



Appendix 2: fauna! lists 

Pygomeles braconnieri Grandidier, 1867 R 
ENDEMIC. SW - Toliara area. Fossorial, in sand. 

Pygomeles petteri Pasteur et Paulian, 1962 

ENDEMIC. NW - Ankarafantsika. Known by types only. 

Voeltzkowia fierinensis (Grandidier, 1869) K 
ENDEMIC. SW - S of Toliara on Fiherenana River. 

Voeltzkowia lineata (Mocquard, 1901) K 

ENDEMIC. S, SE.SW. In SW appears not to occur N of R. 
Fiherenana. Most northerly point Sakaraha (note on habitat 
in Brygoo 1981). 

Voeltzkowia mira Boettger, 1893 K 

ENDEMIC. NW. Mahajanga region. Holotype found in 
rotten trunk of 'Sabra' Palm Hyphaene coriacea , other 
specimens in sand. 



Voeltzkowia petiti (Angel, 1924) 
ENDEMIC. SW - Scrub dunes 
Toliara. Two specimens only. 



at Tsivono, 24 km of 



Voeltzkowia rubrocaudata (Grandidier, 1869) K 
ENDEMIC. SW (Fierin: Toliara). N of Toliara ami the R. 
Fiherenana. Beroboka between Morondava and Belo 
Tsiribihina. 



Cryptoposcincus minimus Mocquard, 1906 

ENDEMIC. 'Madagascar' - locality unknown. Known by 

holotype only. 



Mabuya aureopunctata (Grandidier, 1867) 
ENDEMIC. S, W. In arid scrub. 



nt 



Mabuya betsileana Mocquard, 1906 

ENDEMIC. C - Betafo in Betsileo Prov. Known by type 

only. Validity of taxon uncertain, geographic error possible. 

Mabuya boettgeri Boulenger, 1887 R 
ENDEMIC. C, E-C. 

Mabuya elegans (Peters, 1854) nt 

ENDEMIC. Throughout except C and S-C. One of 
commonest lizards in NW. Along littoral, arid scrub, under 
stones in subdesert areas. Includes in synonymy: M. 
sakalava (Grandidier, 1872), Brygoo 1983, in press. 

Mabuya gravenhorsti (Dumeril et Bibron, 1839) nt 
ENDEMIC. Very common, found throughout, 0-1000m, on 
coastal rocks, scrub, wooded hills. Favoured by extension of 
'savoka' secondary vegetation. 

Mabuya madagascariensis Mocquard, 1908 K 

ENDEMIC. Type specimen 'Madagascar' - no locality, 

recently rediscovered on Tsiafajavona (Ankaratra massif). 

Paracontias brocchii Mocquard, 1894 R. 
ENDEMIC. N - Montagne d'Ambre. 

Paracontias hildebrandti (Peters, 1880) K 
ENDEMIC. NW, Nosy Be. 

Paracontias holomelas (Giinther, 1877) R 

ENDEMIC. E - Anzahamaru, near Mahanoro (terra typica). 



Paracontias milloti Angel, 1949 
ENDEMIC. Nosy Mamoko, 
Known by holotype only. 



Ambariotelo Archipelago. 



Paracontias rothschildi Mocquard, 1905 

ENDEMIC. 'Madagascar' - locality unknown. Known by 

types only. 

Pseudoacontias madagascariensis Barboza du Bocage, 1889 
ENDEMIC. Locality unknown. Known only from Holotype, 
recently (1979) destroyed by fire at the Bocage Museum, 
Lisbon. 



Family CORDYLIDAE (GERRHOSAURINAE) 
Tracheloptychus madagascariensis Peters, 1854 
ENDEMIC. N (Nasatra), E, SE, S, W. 

Tracheloptychus petersi Grandidier, 1869 
ENDEMIC. SW - Morombe, Tsivanoha. 

Zonosaurus aeneus (Grandidier, 1872) 

ENDEMIC. E, W, S-C. Wooded hilly areas, littoral areas. 

Zonosaurus boettgeri Steindachner, 1891 
ENDEMIC. NE, Nosy Be. 

Zonosaurus karsteni (Grandidier, 1869) 
ENDEMIC. W, SW, S-C. 

Zonosaurus laticaudatus (Grandidier, 1869) 
ENDEMIC. NW, E, SE, S, SW, C. 

Zonosaurus madagascariensis (Gray, 1845) 

NW, NE, E coast, SW, Nosy Be, St. Marie; also Glorieuse. 

Littoral, wooded areas to 1000m, including primary forest. 

Zonosaurus maximus Boulenger, 1896 

ENDEMIC. SE, E-C, S-C, C. Forest species, also found 
along rivers and watercourses. Frequent on banks of the 
Faraony. Found above all in dense leaf litter of mango trees 
where it seeks earthworms. One of the largest Malagasy 
lizards, to 67cm. 

Zonosaurus ornatus (Gray, 1845) 

ENDEMIC. NW, N-C, E, SE, SW, S-C, C. On rocks, in 

forest. 'Antsiantsy'. 

Zonosaurus quadrilineatus (Grandidier, 1867) 
ENDEMIC. SW. 

Zonosaurus rufipus (Boettger, 1881) 
ENDEMIC. E, Nosy Be. 

Zonosaurus trilineatus Angel, 1939 
ENDEMIC. Ambovombe. 



•185- 



An environmental profile of Madagascar 

SERPENTES 

NB Status designations and other comments from Domergue 
( in litt. , rec 27.6.83). 

Family TYPHLOPIDAE 

Ramphotyphlops braminus (Daudin, 1803) 
Nossi-be (NW), Ambilobe (NW), Betsako (NW), Toamasina 
(E), Ambatolampy (C), Mandraka forest (C), Ambovombe 
(S); worldwide. 

Typhlops arenarius (Grandidier, 1872) 

ENDEMIC. Morondava, Mahajanga (W), Menabe (W), 

Toliara (W), Andrahomana (S). 

Typhlops decorsei Mocquard, 1901 

ENDEMIC. Ambovombe (type), southwest around Toliara. 

Typhlops grandidieri Mocquard, 1905 
ENDEMIC. "Madagascar". 

Typhlops madagascariensis Boettger, 1877 
ENDEMIC. Nosy Be. 

Typhlops microcephalus Werner, 1909 
ENDEMIC. "Madagascar". 

Typhlops mucronatus Boettger, 1880 
ENDEMIC. ? 

Typhlops ocularis Parker, 1927 

ENDEMIC. Known only by the type from Antongil forest, 

Maroantsetra region (NE). 

Typhlops reuteri Boettger, 1881 
ENDEMIC. Nosy Be. 

Family BOIDAE 
Acrantophis dumerilii Jan, 1860 

ENDEMIC. South and southwest. Still moderately 
common. Localities include: Amboasary (S), Belo (W), 
Mahabo (W), Morondava (W), Andranolava (C), Toliara 
(S). Frequents the humid edges of pools and water-courses. 
•Do'. 

Acrantophis madagascariensis (Dumeril et Bibron, 1844) 
ENDEMIC. West (north of the Tsiribinina), north, 
northeast, east. Still moderately common. More rare than 
Sanzinia . Localities include: Mahajanga (W), Amboasary 
(W), Ste. Marie de Marovoay (S), Nosy Be. Frequents the 
humid edges of pools and water-courses. 'Do' (in general) or 
'Ankoma' (in East). 

Sanzinia madagascariensis (Dumeril et Bibron, 1844) 
ENDEMIC. Occurs throughout Madagascar, but especially 
in the north and the eastern forests. Still moderately 
common. Localities include: Ankafana (E), Mont de 
Francais (E), Southeast of Betsileo, Emininy (E), 
Frandrarazana (E), Mananjary (E), Mandotra (E), 
Toamasina (E), Tampina (E), Androhinaly (S), 
Toliara-Tsihombe-Morondava. The smallest malagasy boa, 
arboreal. 'Manditra' (in east). 

Family COLUBRIDAE 
Alluaudina bellyi Mocquard, 1894 R 

ENDEMIC. Until recently known only by type from Mt 
d'Ambre (Diego-Suarez)(N); four new specimens comprise 
two from Mt d'Ambre, one from Sambava, one from 
Ambatonutatao. A small secretive species. 

Alluaudina mocquardi Angel 1939 R 
ENDEMIC. Known from type only found in 
Ankara cave. 

Dromicodryas bernieri (Dumeril et Bibron, 1854) nt 
ENDEMIC. Widespread, found throughout the island, very 
common. Includes in synonymy: Liopholidophis 

pseudolateralis Guibe, 1956 (fide Domergue, in litt., 
27.06.83). 



Dromicodryas quadrilineatus (Dumeril et Bibron, 1854) nt 
ENDEMIC. Widespread, found throughout the island, less 
common than D. bernieri . 

Geodipsas heimi Angel, 1936 V 

ENDEMIC. Known from type, from Tsihanovoka on the 

river Sahandrata, also the Zafimanirihy area (forests east of 

Ambositra). 

Geodipsas infralineata (Giinther, 1882) V 

ENDEMIC. Occurs on the plateau and in eastern forests, 

localities include: Moramanga (E), Maroantsetra (NE) 

Heteroliodon torquatus Boettger, 1913 R 

ENDEMIC. Until recently known only by the type, from 

Andranohinaly (near Toliara) (W); two new specimens are 

from north of Toliara and Kinkony Forest, south of 

Morondava. 

NB. The genus Ithvcyphus is in course of revision, there 
appear to be several undescribed forms (Domergue, in litt , 
27.06.83). 

Ithycyphus goudoti (Schlegel, 1854) V 

ENDEMIC. Moderately widespread in NE, E and S, also 

known from the east coast. 

Ithycyphus miniatus (Schlegel, 1837) V 

ENDEMIC. Nosy Be (NW), Andrahomana (S), Andranolava 

(C), Fandrazana (E). 'Fandrefiala'. 

Langaha alluaudi Mocquard, 1901 V 

ENDEMIC. Occurs throughout the island in forest areas, 

localities include: Andrahomana (S), Behara (S), Menabe 

(W), Ambovombe (S), Taolanaro (SE), Bemamanga. 

'Fandrefiala'. 

Langaha nasuta Shaw, 1790 V 

ENDEMIC. Occurs throughout the island in forest areas, 
localities include: Nosy Be (NW), Antongil Bay (NE), 
Ambatomainty (O), Ankarafantsika (O), Imerina (C), 
Betroka (C), Befanany (SW), Morondava (E). 

Leioheterodon geavi Mocquard, 1905 nt 

ENDEMIC. Occurs throughout the south, south of the 

Tsiribihina River, localities include: Fiherana plain (type), 

Ambovombe (S), Toliara (SW), Betioky (SW), Androka 

(SW). 

Leioheterodon madagascariensis Dumeril et Bibron, 1854 nt 
ENDEMIC. Rather common and widespread, occurs 
throughout the island but especially the north and east. 
'Menarana'. 

Leioheterodon modestus (Giinther, 1863) nt 
ENDEMIC. Found throughout the island, especially the 
west and north, also the south as far as Antsirabe. 
'Bemavo', 'Le Fompoty'. 

Liophidium apperti Domergue, 1983 K 

ENDEMIC. Known by a single specimen collected in 1968, 
in deciduous forest 7km north of the village of 
Befandriana-sud, 2km east of RN 9 (Toliara-Morombe). 
This forest has now been cleared; only a few baobabs, 'kily' 
and 'sakoas' remain. 

Liophidium rhodogaster (Schlegel, 1837) V 
ENDEMIC. Known from forest regions of the east and 
north, localities include: Antananarivo (C), Ikongo massif 
(SE), Ambatomainty, Beforona (E), Nosy Be. 

Liophidium chabaudi Domergue, 1983 K 
ENDEMIC. Known from three specimens; the holotype from 
Ankorongo (near Toliara airport), the second from a dozen 
km N of Toliara (between Belalanda and Tsongoritela), the 
third from Bevato (south of Morombe). A sub-fossorial 
form, occurring in somewhat swampy and saline areas, 
behind coastal dunes in the littoral zone. 



186- 



Appendix 2: fauna! lists 



Liophidium therezeni Domergue, 1983 K 
ENDEMIC. Known by two specimens collected in 1966 and 
1969; the former (holotype) from Anatelo forest bordering 
the Ankarana massif (Ambilobe sub- prefecture), the latter 
from Antseranana (Antsiranana). 

Liophidium torquatus (Boulenger, 1888) V 
ENDEMIC. In humid forest. Localities include: Antongil 
Bay (NE), Mananjary (E), Anamalazoatra (E), Toamasina 
(E), Ambositra (C), Ambatodratino (C), Ankirika (C), Ste. 
Marie de Marovoay (NW), Taolanaro (SE), also the banks of 
the River Mangoky (SW). 

Liophidium trilineatum Boulenger, 1896 R (K) 
ENDEMIC. Type from SW, also known from 

Tsimanampetsotsa and several new specimens from other 
parts of the south. 

Liophidium vaillanti (Mocquard, 1901) R (K) 
ENDEMIC (?). " Ambovombe (S), Betroka (S-C), 
Maevatanana (NW), (also reported from Reunion, but this is 
extremely improbable). 

Liopholidophis grandidieri Mocquard, 1904 E (K) 
ENDEMIC. Until recently known from the holotype only, 
from the mouth of the River Saint Augustin (SW), three new 
specimens are now known from the eastern forests. 

Liopholidophis lateralis (Dumeril et Bibron, 1854) nt 
ENDEMIC. Known from almost the entire island, absent 
from coastal regions to the south of Toliara. Common on the 
high plateau, frequently found in parks and gardens 
(including Tsimbazaza). 

Liopholidophis pinguis Parker, 1925 I 

ENDEMIC. Antisihanaka (type), also Moramanga and/or 

(?) Alaotra. 

Liopholidophis sexlineatus (Giinther, 1882) nt 
ENDEMIC. Occurs mainly on the east coast and the 
plateau, localities include: Betsileo area (C), Imerina (C), 
Ambatomainty (W), Toamasina (E). A semi-aquatic form, 
abundant in rice cultivation. 

Liopholidophis stumpffi (Boettger, 1881) 

ENDEMIC. Most specimens are from Perinet; six other 
localities are known, mostly along the east coast (including 
on Marojezy), also Nosy Be. A humid forest form, typically 
found in the morning along forest tracks, basking in patches 
of sunlight. 

Liopholidophis thieli Domergue, 1972 

ENDEMIC. A humid forest form, found at several localities 
in the east region, including Perinet (where the type was 
found in fish culture ponds), also on Nosy Mangabe. Eats 
amphibians. 'Menamaso'. 

NB. The genus Lvcodryas is in the course of revision 
(Domergue, in litt., 27 June 1983). 

Lvcodryas arctifasciatus (Dumeril et Bibron, 1854) R 
ENDEMIC. Antananarivo (C), Moramanga (CE), Nosy Be 
(NW). 

Lvcodryas betsileanus (Giinther, 1880) R 

ENDEMIC. Betsileo area (C), also one new specimen from 

Maroantsetra (Antongil Bay). 

Lvcodryas gaimardi (Schlegel, 1837) R 

L.g. gaimardi , Taolanaro (SE), Imerina (C), Nosy Be (NW). 
L.g. granuliceps , Toliara (SW), Fiherana valley, Befaisitra. 
Also Comoro Islands (one specimen ' L.g. comorensis '. 

Lycodrvas guentheri (Boulenger, 1896) R 
ENDEMIC. Ambovombe (S), + SW. 

Lvcodryas inornatus (Boulenger, 1896) R 
ENDEMIC. Only three specimens known, two of unknown 
origin, one from the Didierea forest of Ampotaka (south of 
the Menarandra river in the Beloha-Androy area). 



Lvcodryas maculatus (Giinther, 1858) R 

Known from type, source unknown, and three specimens 

from the Comoros. 

Lvcodryas variabilis (Boulenger, 1896) R 
ENDEMIC. 'Madagascar', no details. 

Madagascarophis colubrinus (Schlegel, 1837) nt 
ENDEMIC. Found throughout the island. Several 

geographic forms. Still fairly common. Found most 
frequently in termitaria. Malagasy name: 'Renivitsika' or 
Mere de Fourmis (= Mother of Ants). 



Mimophis madagascariensis Giinther, 1868 
ENDEMIC. A plateau species. 



nt 



Mimophis mahfalensis (Grandidier, 1867) nt 

ENDEMIC. Widespread, especially the south and 

southwest. Still common. Feeds on snakes. 

Micropisthodon ochraceus Mocquard, 1894 R 
ENDEMIC. Nosy Be, second specimen from unknown 
locality and a few new specimens from Perinet 
(Analamazaotra). Probably arboreal. 

Pararhadinea albignaci Domergue, 1984 K 
ENDEMIC. Known by holotype only, from R.S. 19 
Analamazaotra (Perinet); found in January 1970, dead on 
road after heavy rain, between the village of Perinet and the 
forest station. 

Pararhadinea melanogaster Boettger, 1898 R 

ENDEMIC. Nosy Be, also two new specimens from Perinet 

and Marojezy. A small secretive species. 

Pseudoxyrhopus ambreensis Mocquard, 1894 R 
ENDEMIC. Known only from type, from Montagne 
d'Ambre. 

Pseudoxyrhopus dubius Mocquard, 1904 I 

ENDEMIC. Occurs in the eastern forests, localities include: 

Ikongo (SE), Ambatomainty. Several specimens recorded 

during the last two decades. A nocturnal species, probably 

terrestrial. 

Pseudoxyrhopus heterurus (Jan, 1893) R 

ENDEMIC. Nosy Mangabe (Bloxam in litt. 23.07.86); no 

other locality records or recent records. 

Pseudoxyrhopus imerinae (Giinther, 1888) R 

ENDEMIC. Imerina (C), other specimens from Mt Ibity and 

Mt Tsiafajavony (central plateau). 

Pseudoxyrhopus microps Giinther, 1881 R 

ENDEMIC. Betsileo region, Ankarana. No recent records. 

Pseudoxyrhopus occipitalis Boulenger, 1896 R 
ENDEMIC. Known from type, from SW, not found again. 

Pseudoxyrhopus quinquelineatus (Giinther, 1881) V 
ENDEMIC. Betsileo region, east coast, northwest, one 
specimen from the Mahafaly Plateau (S). 

Pseudoxyrhopus tritaeniatus Mocquard 1894 I 
ENDEMIC. Known only from type, from Betsileo region. 



Also two widespread sea snakes (Hydrophiidae), Pelamis 
platurus and Enhydrina schistosa , are recorded in coastal 
waters. 



Two nominal species, Compsophis albiventris Mocquard, 1894, 
from the Montagne d'Ambre, and Geodipsas boulengeri 
(Peracca, 1892) from near Androngoloaka, are probably based 
on mis-identified Geodipsas heimi (Domergue, in litt., 27 June 



1983): 



•187- 



An environmental profile of Madagascar 

MADAGASCAR AMPHIBIA 

(T) = Type locality. Geographic distribution of several species 
apparently not known in full, or published data not available. 
References are provided in Part V.3. Blommers-Schlosser has 
been abbreviated to B.-S. 

Family HYPEROLIIDAE 

*Note: there are probably no true Hyperolius species in 
Madagascar; species listed by Guibe, 1978, appear to be 
assignable to either Heterixalus (Hyperoliidae) or Boophis 
(Rhacophoridae) (B.-S., 1979 II, and 1982). 

'Hyperolius' arnoulti Guibe, 1975 

Manompana (T), east coast, opposite Nosy Borah. Probably 

a Heterixalus species, B.-S., 1982. 

' Hyperolius' friedrichsi Ahl, 1930 

'Madagascar' (T). Probably a Boophis species, B.-S., 1982. 

' Hyperolius' nossibeensis Ahl, 1930 R 
Nosy Be (T). 

Heterixalus alboguttatus (Boulenger, 1882) 

South-east Betsileo (T), also dunes at Mananjary (east 

coast) and Ranomafana (highroad R.N. 25). 

Heterixalus betsileo (O. Boettger, 1881) 

Imerina (T), centre of Madagascar, Andanolava. A very 
common sedge frog, in savannah like areas and cleared forest 
in the central plateau and west. Synonym: H. renifer , fide 
B.-S., 1982. 

Heterixalus boettgeri (Mocquard, 1902) 

? Removed from synonymy of H. tricolor , B.-S., 1982. 

Heterixalus madgascariensis (Dum. et Bib., 1841) 
'Madagascar' (T), Salavaratse. 

Heterixalus mocquardi (O. Boettger, 1913) 

Taolanaro (T), another specimen collected on leaf in 

Taolanaro. 



Heterixalus rutenbergi O. Boettger, 1881 

Imerina (T), also collected in grassland 20 km west 

Antananarivo. Removed from Hyperolius . B.-S., 1982. 



of 



Heterixalus tricolor (O. Boettger, 1881) 

Nosy Be (T), Isaka, and east coast. A very common sedge 
frog, in dunes, savannah-like areas and cleared forest on the 
east coast. 

Family MICROHYLIDAE: Subfamily COPHYLINAE 

(endemic). 

Anodonthvla boulengeri F. Miiller, 1892 

Madagascar, Anevoka. Known from the East Region, the 
Anosyennes chain including Andohahela. Also collected near 
Fenoarivo Atsinanana, Perinet, Ranomafana (Fianarantsoa), 
Foulpointe and 25km north of Toamasina. Many of these 
were collected in hollow leafstalks of young Ravenala , which 
often harboured antnests; ants were the only food items 
recovered from these specimens. Larval development similar 
to other Cophylines. 

Anodonthvla montana Angel. 1925 

Andringitra mountains (T), abundant at altitude. 

Anodonthvla rouxae Guibe, 1974 K 

Anosyenne chain (T), in bamboos (poss. type loc. only). 

Cophvla phvllodactvla O. Boettger, 1880 K 

Nosy Be (T), known from Nosy Be, Nosy Komba and Mt. 

d'Ambre. 

Madeca8sophrvne truebae Guibe, 1974 K 

Anosyenne chain (T), known only from the type locality. 

Mantipus bipunctatus Guibe, 1974 

Forest of Fivahona (T), Andringitra mountains (prob. type 

loc. only). 



Mantipus guentherpetersi Guibe, 1974 

Tsaratanana mountains (T), 2600m (prob. type loc. only). 

Mantipus inguinalis (Boulenger, 1882) 

East Betsileo (T), known from east and north-east 

Madagascar, including the Andringitra mountains. 

Mantipus laevipes (Mocquard, 1895) 
Montagne d'Ambre (T), type locality only. 

Mantipus minutus Guibe, 1975 

Marojejy mountains (T), (prob. type loc. only). 

Mantipus serratopalpebrosus Guibe, 1975 
Marojejy mountains (T) (prob. type loc. only). 

Paracophvla tuberculata Millot & Guibe, 1951 R 

Perinet Forest (T); where reported in leaf axils of Pandanus 

and Crinium firmifolium , 

* Platyhyla alticola (Guibe, 1974) 

Tsaratanana mountains (T) (poss. type loc. only). *Guibe 

considers Platyhyla a synonym of Platypelis . 

Platypelis barbouri Noble, 1940 
Fanovana Forest (T). 

Platypelis cowani Boulenger, 1882 
East Betsileo (T). 

Platypelis grandis (Boulenger, 1889) 

Madagascar (Taolanaro, Sakana), known from East Region 
and mountain massif?. Recently collected near Anjozorobe 
(alt 1300m, 60km north of Manjakandriana); near Perinet 
(alt. 900m); and near Ampasinambo (alt 500m, 55km west of 
Nosy Varika). Specimens were collected in pairs in water 
holes in tree trunks (Ficus sp.) and a water filled axil of 
Ravenala . A resting male was collected beneath rotting 
wood in the forest. Calling males, always hidden in tree 
holes, are heard at night from September to October. Eggs 
are laid in water inside these holes, the female leaves the nest 
several days after oviposition but the male remains until the 
brood have developed into froglets. It is possible that the 
nursing father provides the larvae with fungicidal protection. 

Platypelis milloti Guibe, 1950 R 

Nosy Be (T), known only from Lokobe Forest on Nosy Be 

(where it lives in leaf axils of Typhonodorum lyndleianum ). 

Platypelis pollicaris Boulenger, 1888 

Madagascar, known from East and Centre Regions and 

Tsaratanana mountains. Collected by B.-S. near Perinet 

and near Tampoketsa d'Ankazobe in leaf axils. May feed on 

ants. 



Platypelis tsaratananaensis Guibe, 1974 

Tsaratanana mountains (T) in bamboo forest 2600m, (prob. 

type loc only). 

Platypelis tuberculata (Ahl, 1929) 
North-west Madagascar (T). 

Platypelis tuberifera (Methuen, 1920 (1919)) 
Ambatoharanana (T). Later collected at Perinet, Fenoarivo 
Atsinanana and Foulpointe. All specimens were found in 
water-filled axils of Pandanus . Food items recovered include 
ants. Larval development likely to be similar to other 
Cophylinae described. 

Plethodontohyla alluaudi (Mocquard, 1901) 
Taolanaro (T). 

Plethodontohyla brevipes Boulenger, 1882 
East Betsileo (T), a pair were collected by Peyrieras under 
decaying wood in forest near Ampasinambo. The stomachs 
contained beetles. 

Plethodontohyla coudreaui Angel, 1938 
Betampona Reserve (T). 



188- 



Appendix 2: faunal lists 



Plethodontohyla laevis (O. Boettger, 1913) 
Sakana (T), (Tsihanovoha Forest); a specimen was collected 
by Peyrieras near Tampoketsa d'Ankazobe (alt. 1600m) from 
a burrow. 

Plethodontohyla notosticta (GUnther, 1877) 
Mahanoro and Anzahamara, known from forest areas in East 
Region and at low altitude in the Andringitra and Marojejy 
mountains. Collected by B.-S. near Ranomafana, Fenoarivo 
Atsinanana and Foulpointe. Larval development similar to 
that of Platyhyla grandis . 

Plethodontohyla ocellata Noble & Parker, 1926 
Antsihanaka (T), known from the East Region and Marojejy 
mountains. One gravid female collected by Peyrieras under 
decaying wood in forest near Ampasinambo (alt. 500m, 55km 
west of Nosy Varika). The stomach contained a large beetle. 

Plethodontohyla tuberata (Peters, 1883) 

'Madagascar' (T), (Manjakatompo). Found on the high 
plateaus, common at 1500-2000m in the Andringitra and 
Ankaratra mountains. A fossorial form inhabiting forest 
humus. 'Sahondokoro'. 

Rhombophryne testudo O. Boettger, 1880 
Nosy Be, Marojejy and Reunion. 

Stumpffia grandis Guib6, 1974 

Marojejy mountains (T) 1300m, (possibly type locality only). 

Stumpffia psologlossa Boettger, 1881 

Nosy Be (T). Known from East Region, also Sambirano and 

Tsaratanana and Marojejy mountains. 

Stumpffia roseifemoralis Guibe, 1974 

Only known from Marojejy mountains (T), 1300m. 

Stumpffia tridactvla Guib£, 1975 
Marojejy mountains (T). 

Family MICROHYLIDAE: Subfamily DYSCOPHINAE 

( Dvscophus is endemic genus). 
Dyscophus antongili Grandidier, 1877 I 

Antongil Bay (T), localised at Antongil Bay (Moroantsetra, 
Foizana) and south of Toamasina (Andevoranto). A well 
known species, dull red-orange in colour, growing to large 
size (c 9cm). 'Sahongongona', derived from the distinctive 
call. 



Dyscophus guineti (Grandidier, 1875) 

Sambava (T), known from Soalala (north-west) 

(north-east) and Antsihanaka (centre). 



Sambava 



Dyscophus insularis Grandidier, 1872 

Antsouhy (T) near Trabouzy, (south-west Madagascar, 

Belo, Soalala). Reported from the west and south-west, 

specimens recently received from the Marojejy massif (300 

m), others from Ankarafantsika. Includes in synonymy: D. 

quinquelineatus . 

Family MICROHYLIDAE: Subfamily SCAPHIOPHRYNINAE 
(endemic) (NB., sometimes placed in RANIDAE) 
Pseudohemisus calcaratus (Mocquard, 1895) 
Madagascar, south-west, Ambongo, West Region. 

Paeudohemisus granulosus Guibe, 1952 

Andranoboka, Mahajamba Bay north of Mahajanga, also 
near Ampijora in the Ankarafantsika forest, west 
Madagascar. Adult specimens collected on the forest floor, 
tadpoles have been found in shallow temporary pool in the 
shade of large mango tree in January. 

Pseudohemisus madagascariensis (Boulenger, 1882) 
East Betsileo, Andringitra mountains. One of the most 
strikingly coloured Malagasy frogs; bluish to olive green with 
sinuous brown dorsal stripes. 

Pseudohemisus pustulosus Angel et Guibe, 1945 
Madagascar, Ankaratra mountains. 



Scaphiophrvne marmorata Boulenger, 1882 

East Betsileo, Foizana, forest species in East Region. 

Family MICROHYLIDAE: Subfamily MICROHYLINAE 
Microhyla palmata Guibe, 1974 
Ambana (T) (90m), Anosyenne chain. 

Family RANIDAE 
Ttychadena mascareniensis (Dum. & Bib., 1842) 
NON-ENDEMIC, occurs elsewhere in East Africa. 
Bourbon (T); very common in all regions, notably 
modified habitats (e.g. rice fields). 



He 



* Rana tigrina Daudin, 1803 
NON-ENDEMIC, an asiatic species 
north-west coastal Madagascar from the 
Increasingly invading the high plateau. 



introduced to 
Indian region. 



Tomopterna labrosa Cope, 1868 

Madagascar (T); A fossorial form, absent from the eastern 
forests. Known from the west (regions of Mahajanga, Belo, 
Soalala, Antseranana) and the south (around Toliara and 
Taolanaro). Has been found on the foothills of the 
Ankaratra range. 

Family RANIDAE: Subfamily MANTELLINAE (endemic). 
Laurentomantis horrida (O. Boettger, 1880) R 
Nosy Bi (T). Found since in Marojejy mountains. 

Laurentomantis malagasia (Methuen & Hewitt, 1913) R 
Folohy (T). 

Laurentomantis ventrimaculata (Angel, 1935) R 
Isaka Ivondro (T), Taolanaro area. 

N.B. The above three species formerly assigned to the genus 
Trachymantis . 

Mantella aurantiaca Mocquard, 1900 V 

Madagascar, only known from the forest of Perinet 
(Antaniditra). Occurs in distinctive swamp rain forest with 
Pandanus species. The species has been collected for many 
years for zoos and terrarium- keepers (B.-S., in litt. . Jan. 
1983). Orange-red, probably poisonous, active during the 
day when great numbers may be seen jumping on the forest 
floor. (See Arnoult 1966 and Oostveen 1978 for ecology, also 
B.-S., 1975.) 

Mantella betsileo (Grandidier, 1872) 

Betsileo (T), (Nosy Be), forested regions of the east. 

(Widespread; map of localities in Busse 1981). 

Mantella laevigata Methuen it. Hewitt, 1913 V 

Folohy (T), east Madagascar. (NB., map in Busse 1981 

indicates Nosy Mangabe as only site). 

Mantella madagascariensis (Grandidier, 1872) 

Incl. M. cowani Boulenger, 1882 (part) plus M. pulchra 

Parker, 1925. Three ssp., widespread (Busse 1981). 



Mantidactylus acuticeps Ahl, 1929 
'Central Madagascar' (T); type lost. Found at 
(highroad R.N. 2) alt. 900-1100m, in the evening i 
thicket in forest, far from water. 



Perinet 
i dense 



Mantidactylus aerumnalJB (Peracca, 1893) 

Andrangoloaka (T)(type lost); Andringitra. Known from 
central region and Andringitra and Ankaratra mtns. Found 
in forest beside brooks and in more open areas beneath 
shrubs near forest. Tadpoles found in quiet parts of the 
same streams. 

Mantidactylus aglavei (Methuen et Hewitt, 1913) 
Anamalazotra (T); Moramanga, Andringitra and 
Moramanga forest, East Region. Tree frogs. Males found 
calling in trees in the evening. Tadpoles found in a shallow 
brook in open land near forest. 



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An environmental profile of Madagascar 



Mantidactvlus albofrenatus (F. Miiller, 1892) 
'Madagascar' (T); known from the eastern forests, 
Foulpointe, also found in the West Region (forest of 
Mohajeby, region of Morafenobe). Found along a brook in 
the forest during the day, the males calling in dense 
vegetation. Surface-feeding tadpoles found in the same 
brook adhere to the surface by the tail in dense aquatic 
vegetation. 

Mantidactvlus alutus (Peracca, 1893) 

Andrangoloaka (T); East Region, Central Region. Collected 
in and along rice fields and pools. Tadpoles were in nearby 
ditches and pools. Also found in small brooks in open land. 

Mantidactylus ambohimitombi Boulenger 1919 
Ambohimitombi forest (T); East Region. A specimen was 
found on the bank of a clear brook with a stony bottom in 
forest. 

Mantidactvlus argenteus Methuen, 1920 (19191 R 
Folohy, east Madagascar (T); Andringitra mountains. 

Mantidactylus asper (Boulenger, 1882) 

East Betsileo (T); Known from the eastern region and the 
Marojejy, Andringitra and Tsaratanana mountains. Occurs 
in well-developed forest and dense shrubwood. Adults climb 
like tree-frogs, but are also found on the ground. Most 
active at dusk and in early morning. Eggs deposited singly 
or in pairs; direct development. Reproductive season 
probably limited to a few months (end Oct. - early Jan.) in 
the wettest period of the year (the egg needs a permanently 
damp atmosphere and the tiny froglet a choice of small prey, 
these conditions are only met in the rainy midsummer). 

Mantidactvlus bertini (Guibe, 1947) 

Isaka Ivondro (T); known from East Region and Anosyenne 

chain. 

Mantidactvlus betsileanus (Boulenger 1882) 
Betsileo (T); west Madagascar. Common in all areas except 
the south. Common in boggy places in open degraded forest, 
sometimes mixed with M. biporus . Tadpoles found in 
shallow pools, often with those of M. liber . Egg mass found 
fixed to a leaf on ground in wet grassland. 

Mantidactylus bicalcaratus (Boettger, 1913) 
Nosy Borah (T). Apart from type locality the species is 
known from the Marojejy mountains and Taolanaro. Also 
25km north of Toamasina, sea level; Foulpointe (60km north 
of Toamasina), sea level; midway between Foulpointe and 
Fenoarivo Atsinanana, sea level; Ambila-Lemaitso (near 
Brickaville), alt. 7m; Mananjary, alt. 13m. Very abundant 
on the east coast. Adults found in the leaf axils and leaves of 
PandanuB dauphinensis in dunes near the sea and around 
lagoons, and in Typhonodorum lindlevanum in coastal 
swamps. Tadpoles in leaf axils. 

Mantidactylus biporus (Boulenger, 1889) 

'Madagascar' (T). A forest species from the East Region. 
Occurs in small shallow muddy pools, slow flowing water in 
open land adjacent to forest, gutters of roads, shallow water 
between Rapphia palms and small pools along streams. 

Mantidactvlus blanci Guibe, 1974 

Andringitra (T); Known only from Andringitra mtns. 

Mantidactylus blommersae (Guibe, 1975) 

Moramanga and Perinet forests (T). Known from type 
localities, also Perinet (highroad R.N. 2) alt. 900-1100m, 
Mandraka Valley (highroad R.N. 2 at km 69), rivulet 
Vokanatezandava and adjacent ponds, nursery garden of the 
city of Antananarivo, alt. 1200m., between Ranomafana and 
Ifanadiana (highroad R.N. 25) alt. 800m. Adults collected in 
November - April at breeding sites near sunlit pools (often 
temporary) 1cm - lm deep, together with egg masses 
(attached to leaves a few feet above the water) and 
tadpoles. At other times adults were captured on the ground 
in forest. 



Mantidactvlus brevipalmatus E. Ahl. 1929 
North-west Madagascar (T) - type lost. There is some 
doubt over the validity of this species since the type has 
been lost. 

Mantidactvlus boulengeri Methuen, 1920 (1919) 
Folohy (T), (Fort Carnot). Common in forests in the East 
Region. Occurs on the forest floor in dense vegetation 
Males do not form choruses when calling but are widely 
dispersed. This may indicate direct development of the egg 
and possibly parental care. They are particularly vulnerable 
to exposure to the sun. 

Mantidactvlus bourgati Guibe, 1974. 

Andringitra (T). Known only from the type region, 
including:- Boby basin (2500m), Marositry stream (2000m), 
Amparabatosoa plateau (2100m), Antanfotsy village 
(1450m), Ampanasana River; Ambohambatomanara col 
(2100m); Ambalamaisinjo plateau: Riambouy River 
(2000m); Akiseasea (1500m); Ambalamarovandana (1500m). 

Mantidactvlus curtus (Boulenger, 1882) 

East Betsileo (T). Known from the East Region and 
mountain areas. Adults are found in and along streams, 
mainly in open areas but sometimes in forest. Tadpoles are 
found in the lentic sidepools of the same streams. 



Mantidactvlus decarvi (Angel, 1930) 

South Midongy and Andringitra mtn., Befotaka. 

from the East Region and mountain areas. 



Known 



Mantidactylus depressiceps (Boulenger, 1882) 
Betsileo (T); Sahembendrana, Akkoraka. Known from the 
eastern region. Rests during the day and the dry cool season 
on leaf axils of Pandanus , Typhonodorum lindlevanum , 
Musa and tree ferns. Egg masses are attached to leaves 1-2 
metres above pools. 

Mantidactvlus domerguei (Guibe, 1974) R 
Andringitra (T). Also found at Manjakotompo forest station 
(near Ambatolampy), Ankaratra mountains, alt 
1800-2400m. Ground dwelling. Found on forest floor near 
temporary pools. 

Mantidactylus elegans (Guibe, 1974) 

Massif de 1 Andringitra (T); known from the Andringitra and 

Tsaratanana mountains. 



( Mantidactylus elegans (Guibe, 1974)) 

* NB., replacement name: Mantidactvlus guibei , 

press. Anosyenne chain (T). 



B.-S. 



Mantidactvlus eiselti Guibe, 1975 R 

Perinet Forest (T); known only from type locality. Calling 
males found during the day in brushwood in hillside forest, 
sitting on small branches just above the ground; heard 
calling only during the rainy season, always singly and 
metres apart, far from open water. Particularly vulnerable 
to exposure to the sun. Wide dispersal of calling males and 
distance from water may indicate direct development of the 
eggs and possibly parental care. 

Mantidactylus femoralis (Boulenger, 1882) 
East Betsileo (T), (Mt. d' Ambre); region de Rogez, 
Isaka-Ivondro. Common in the eastern forests. Always 
found on the ground or in shrubs, along clear forest brooks, 
tadpoles found in side pools. 

Mantidactvlus flavicrus (Boulenger, 1889) 

Madagascar (T); known from the eastern regions, Sambirano 

and Montagne d'Ambre. 

Mantidactvlus flavobrunneus B.-S., 1979 
On the road from Moramanga to Anosibe at km 25, alt 
900m.; Perinet (highroad R.N. 2) alt. 900-1100m. Rainforest 
on eastern escarpment. Found in water in Pandanus axils. 



Mantidactvlus glandulosus Methuen & Hewitt, 1913 R 
Folohy (T); known only from type locality, type lost. 
M. pseudasper is probably a synonym, (B.-S., in litt). 



NB. 



•190- 






Appendix 2: faunal lists 



Mantidactylua grandidieri Mocquard, 1895 

East coast (T). Collected in and along brooks in open land 

next to forest. Ground dwelling. Tadpoles unknown. 

Mantidactvlus grandisonae Guibe, 1974 R 

Ambana (T), Anosyenne chain; known only from type 

locality, alt. 1000m, low altitude forest. 

Mantidactvlus guttulatus (Boulenger, 1881) 

South-east Betsileo (T); East Madagascar, Ikongo forest. 

Eastern region. 

Mantidactvlus granulatus (O. Boettger, 1881) 
Nosy Be (T); Found in the eastern forests, Montagne 
d'Ambre, Tsaratanana mountains. Also known from 
Mayotte, Comores. 

Mantidactvlus inaudax (Peracca, 1893) 
Andrangoloaka (T); East Region. 

Mantidactvlus klemmeri (Guibe, 1974) R 

Marojejy mountains (T); known only from type locality. 

Mantidactvlus liber (Peracca, 1893) 

Widely distributed in the central highlands, at 800-1300m; 
this area is characterised by medium altitude rainforest, now 
(1975) confined to isolated patches and a fringe on the steep 
escarpments on the east side of the highlands. Most of the 
collections were made in more or less degraded forest of this 
type. In Tampoketsa d'Ankazobe some of the last vestiges of 
high altitude forest are preserved. M. liber was found in no 
other region. Localities: Andrangoloaka, near 

Manjakandriana, Perinet, Itremo, along the road from 
Ranomafana to Ifanadiana at 5km, along the road from 
Moramanga to Anosibe at 25km, Perinet at an altitude of 
900- 1100m, Mandrake valley along the road from 
Antananarivo to Moramanga at 69km, Anjozorobe alt. 
1300m, and Tampoketsa d' Ankazobe alt. 1600m. Annual 
rainfall 1500- 2000mm, falling almost entirely in the hot 
season, between the end of October - beginning of December 
until the end of March - mid May. The aridity of the dry 
season is moderated by abundant dew formation, 
condensation and frequent fogs. An arboreal frog, inactive 
during the day. Rests in axils of the larger Pandanus , 
Typhonodorum lindleyanum (Araceae), Banana and 
Ravenala madagascariensis (Musaceae), as well as some 
Amarillidaceae ( Crinium firmifolium ), palms f Rapphia ) and 
arborescent ferns. The axils nearly always contain some 
water, even in the dry season. Specimens are found in 
quantity when resting sites near water, together with M. 
methueni and M. pulcher . Becomes active at dusk, feeds 
primarily on small insects such as mosquitoes and flies. 
Apparently breeds from November to May. Exhibits 
unusual mating behaviour which may extend to other 
members of the genus. Eggs are deposited in a gelatinous 
mass on leaves overhanging shady shallow pools, in which 
the hatching larvae develop. 

Mantidactylus lugubris (A. Pumeril, 1853) 

'Madagascar' (T); Betsileo. Common in the eastern region 
and the mountain ranges (Andringitra, Marojejy, Montagne 
d'Ambre). Always found on banks of faster flowing currents, 
or on boulders in rapids. 

Mantidactylus luteus Methuen & Hewitt, 1913 

Folohy, east Madagascar (T). Forest species from the 

eastern region. Found on the forest floor, probably also 

climbs. 

Mantidactvlus madecassus (Millot & Guibe, 1950) 
Andringitra (T). Described from the Cirque Boby in the 
Andringitra massif, has been found since at the same site. 

Mantidactylus majori Boulenger, 1896 

Ivohimanita (T). A forest form from the eastern region. 

Mantidactvlus microtvmpanum Angel, 1935 
Isaka-Ivondro (T). Known form the eastern region (Isaka 
forest, Taolanaro region) and has recently been found in the 
Andringitra massif and the Anosyennes mountains. 



Mantidactvlus opiparis (Peracca, 1893) 

Andrangoloaka (T), near Moramanga. Also Perinet 
(highroad R.N. 2 at km 142), alt 900-1100m, Mandraka 
Valley (highroad R.N. 2 at km 69), rivulet Vokanatezandava 
and adjacent ponds, nursery garden of the city of 
Antananarivo, Tampoketsa d'Ankazobe, forest station alt. 
1600m. Collected in or near forest. Adults jump strongly. 
Tadpoles collected among debris in quiet corners of streams. 

Mantidactylus pauliani Guibe, 1974 

Ankaratra massif:- Nosiarivo & Betay forest, Manjakotompo 
(a forest station nr. Ambatolampy), alt 2200 m, under 
boulders in rapids. So far only known from the type region. 

Mantidactylus peraccae (Boulenger, 1898) R 
Ivohimanita (T); also at Tampoketsa d'Ankazobe, forest 
station, alt 1600m; captured from the axil of Pandanus . 
Related to M. depressiceps . Tree frogs, rest in leaf axils. 
Not observed in open water. 

Mantidactylus pliciferus (Boulenger, 1882) 
East Betsileo (T). Common in mountain ranges. 

Mantidactvlus pulcher (Boulenger, 1882) 

Betsileo (T); Andrangoloaka, Ambila & Lake Aloatra, 
Perinet Forest, Itremo. Very common in all forest regions. 
Inhabits leaf axils. Eggs deposited in one mass above the 
water, in leaf axils. 

Mantidactylus punctatus B.-S., 1979 R 

All specimens collected in axils of Pandanus at Tampoketsa 
d' Ankazobe, forest station alt. 1600m in the relict forest in 
the gulleys of the Tampoketsa. 

Mantidactylus pseudoasper Guibe, 1974 R 
Massif du Marojejy (T). Found in low altitude forest 
(300m). NB., probably a synonym of M. glandulosus , B.-S., 
in litt. 

Mantidactylus redimitus (Boulenger, 1889) 

'Madagascar' (T). Known from the eastern region and the 

massifs of Andringitra, Marojejy and the Anosyenne chain. 

Mantidactylus tornieri (Ahl, 1928) 

Anhoraka, Sahambendrana, central Madagascar, also 
Pennet (highroad R.N. 2 at km 142), alt. 900-1100m, 
Foulpointe (60km north of Toamasina), sea level. Use axils 
of plants such as Ravenala and Typhonodorum 
lindleyanum . In the evening they emerge onto the leaves. 
Egg masses are attached to leaves 30cm - 3m above 
permanent pools. 

Mantidactvlus tricinctus (Guibe, 1947) 

Befotaka and Vondrozo. Known from the eastern forest, in 

the Andringitra mountains and the Anosyenne chain. 

Mantidactvlus ulcerosus (Boettger, 1880) 

Nosy Be (T); Montagne d'Ambre, Akkoraka. A very 
common species in all forest areas. Found in shallow pools 
and slow running water in marshy land or forest. 

Mantidactylus webbi (Grandison, 1953) R 

Nosy Mangabe, Antongil Bay (T). Two individuals from the 

Farankariana forest station have been assigned to this 

species. 

Mantidactvlus wittei Guibe, 1974 

Surroundings of d'Ambanja (T); also Nosy Be, Ampijora, 
Ankarafantsika forest and other western sites. Adults found 
on the forest floor, and in low vegetation near temporary 
pools. Egg masses attached to leaves overhanging water. 
Ground dwelling. 

Family RHACOPHORIDAE 
Aglyptodactvlus madagascariensis (A. Dumeril, 1853) 
'Madagascar' (T); Anzahamaru, north-west Madagascar. 
Very common, found in all forested regions. Ground 
dwelling. B.-S. (1979a) excludes this monotypic genus from 
the Mantellinae. 



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An environmental profile of Madagascar 

Boophis albilabris (Boulenger, 1888) R 

East Imerina (T). Known from the eastern forests. 

Boophis brygooi (Guibe, 1974) 

Andringitra mountains (T); known only from type region. 
An altitude form, appears very common in the Andringitra 
mountains, where it was collected in abundance as adults 
and at various stages of metamorphosis in Nov, Dec, and Jan. 

Boophis callichromus (Ahl, 1928) 
North-west Madagascar, central Madagascar. 

Boophis difficilis (Boettger, 1892) 

Foizana, east Madagascar (T), also Perinet. Never heard or 
seen during daytime. Males were found on rainy evenings in 
forest, calling in vegetation beside brooks. 

Boophis erythrodactylus (Guibe, 1953) 

Forest of Mahajeby (T), cloBe to Morafenobe, west 
Madagascar. Also collected in Mandraka Valley, 

Manjakotompo forest station, Ankaratra Mts, and near 
Perinet. Males call from leaves of shrubs and trees alongside 
rapids, in the evening. 

Boophis goudoti Tschudi, 1838 

'Madagascar' (T). Geographically variable, distributed over 
the whole island. Found in or near stagnant or slow running 
water in forests and ricefields. Pairs in axillary amplexus 
found in August. Tadpoles found in slow running water and 
adjacent pools. Feed on large prey e.g. grasshoppers, moth 
and beetle larvae. Also found in trees although in general a 
poor climber. Eggs are deposited in clumps of 30, attached 
to rocks in water. Reproduction starts at the end of 
November. Valued as a dietary item. 'Sahabakaka'. 

Boophis granulosus (Guibe, 1975) 

Moramanga Forest (T). Young and tadpoles have been 
collected near Perinet, at 900m and 1100m altitude. Adults 
found on leaves of shrubs and trees around a pool, near 
forest; tadpoles found in same pool. 

Boophis hillenii B.-S., 1979 

Near Perinet (T); also near Ranomafana. Males found 
calling during a rainy night in shrubbery around temporary 
pool in forest. Tadpoles found in a temporary pool. 
Sympatric with B. granulosus and B. idae . 

Boophis hyloides (E. Ahl. 1929) 

Central Madagascar (T). Known only from type region. 

Boophis idae (Steindachner, 1867) 

Madagascar, (east Betsileo, Fianarantsoa). Known from East 
Region, Also Mandraka Valley, and near Perinet. Males 
were heard calling in October and November in the evening, 
near stagnant sunlit pools. 

Boophis laurenti Guibe, 1947 

Andringitra (T), Cirque Boby. Known only from Andringitra 

mountains. Uncommon. 

Boophis leucomaculatus (Guibe, 1975) R 

Nosy Mangabe, Antongil Bay. Known only from type 

specimen. 

Boophis luteus (Boulenger, 1882) 

Ankafana, Betsileo, (Moramanga, Antsihanaka). Not rare in 
forests in the East Region. During rainy nights males are 
heard calling from the leaves of trees & shrubs beside rapids 
in the forest. Tadpoles occur in swiftly flowing waters. 

Boophis rrmdagttscariensis (Peters, 1874) 

Madagascar, (Nosy Be, Akkoraka), East Region. Males call 
in the evening in low vegetation less than lm above small 
shallow muddy streams. During the day occasionally found 
in leaf axils of large plants. A true forest species. 

Boophis majori (Boulenger, 1896) 

Ambohimitombi forest (T). A forest species of the East 

Region. 



Boophis mandraka B.-S., 1979 

Mandraka Valley (T), probably type locality only. Altitude 
1200m. Males were caught calling in shrubbery alongside 
rapids, on rainy nights. Tadpoles found in flowing water. 

Boophis microtis (Guibe, 1974) R 

Anosyenne chain (T). Known only from type locality. 

Boophis microtympanum (Boettger, 1881) 
Imerina (T), east Betsileo, known from mountain areas - 
Ankaratra, Andringitra. Collected in clear mountain brooks 
with stony bottoms, in wooded country with ericoid bushes. 
Jumps and climbs poorly. Tadpoles and eggs found in the 
same brooks as the adults. Eggs (clutch of about 100) are 
attached to a twig in fast flowing stream. Axillary amplexus. 

Boophis miniatus (Mocquard, 1902) 

Forest between Isaka and valley of Ambobo near Taolanaro, 
south Madagascar. Also near Perinet. Alt. 900m. Not 
observed during the day. Males found calling in the evening 
in the vegetation beside forest brooks. 

Boophis opisthodon (Boulenger, 1888) 

'Madagascar' (T); forest areas in East Region. A large 
species, probably breeds in temporary pools. Males found 
calling in shrubbery 30-50cm above shallow pools in 
February (these pools had disappeared by July). Calling 
males were around 10m apart. 

Boophis pauliani (Guibe, 1953) 
Forest of Moramanga and Perinet. 

Boophis rappiodes (Ahl, 1928) 

Sahambendrana (T); also near Perinet, Mandraka Valley, 

Moromanga-Anosibe road. Stream breeding. 

Boophis reticulatus B.-S., 1979 

Near Perinet (T), probably type locality only. Males were 
found on rainy evenings, calling on leaves of shrubs and trees 
by running water in open woodland. 



Boophis rhodoscelis (Boulenger, 1882) 

East Betsileo (T), (Andrangoloaka, 

Madagascar), East Region. 



north-west 



Boophis tephraeomystax (A. Dumeril, 1853) 
Madagascar. Very common in forest areas in the East 
Region. Common in all coastal areas. Probably the only 
Boophis sp. occurring in the south-west (the driest part of 
island) where it is found in the irrigated area around 
Toliara. A secretive species, sheltering in leaf axils of plants 
during the dry season. Has the greatest tolerance of drought 
and heat of all Boophis spp., but is also very prominent in 
the humid eastern forests. Tadpoles are found in sunlit 
temporary rainpools with abundant vegetation. 

Boophis untersteini (Ahl, 1928) 

Central Madagascar (T), also north-west Madagascar, 
Mandraka valley, near Perinet, forest station, Tampoketsa 
d'Ankasobe and the road from Moramanga to Anosibe. 
Tadpoles collected in slow running water near forest. 

Boophis viridis B.-S., 1979 

Near Perinet (T), probably type locality only. Males were 
found on a rainy evening, calling on leaves of shrubs and 
trees, beside running water, in forest. 

Boophis williamsi (Guibe, 1974) 

Ambohimirandana (T), Ankaratra mountains. Known only 
from type localities. Also Manjakotompo forest station, 
Ankaratra mounatins altitude 2200 m. Tadpoles were 
collected in clear mountainous brooks with stony bottoms in 
wooded country. Development may take 2 years as in other 
montane spp. 



•192- 






Appendix 2: fauna! lists 



ENDEMIC FISHES OF MADAGASCAR 

* = Freshwater species 

A. ENDEMIC SPECIES 

Family ARIIDAE 
* Ancharius brevibarbis Boulenger, 1911. 

Found in the eastern coastal region and rivers at low 
altitude, the type specimen comes from Ambohimanga. 
Particularly common in rivulets of the eastern escarpment, 
especially around Fort Carnot, Ifanadiana and 
Ambohimanga du Sud. Prefers warm waters. 

* Ancharius fuscus Steindacher, 1880. 

Found in the eastern coastal region and rivers at low altitude 
preferring small riffles and rocky zones. Type specimen 
found at Tohizona. There are also records from Fenoarivo 
Atsinanana, the rivers Vohitra and Rianila near Brickaville, 
River Ranafotsy near Toamasina, Ambanambalo and 
Tohizona (Baie d'Antongil). Little is known about 
reproduction or the ecology of this fish. Easily captured by 
line, the flesh, containing little fat, is widely acclaimed. 

Arius madagascariensis Vaillant, 1894. 

Found more or less throughout the coastal regions, but 
particularly abundant in the lakes and rivers of the west. 
Type locality is Morondava R., W. Madagascar, other 
records being Lake Kinkony and St. Augustin, near Tulear. 
Numerous in lakes and rivers of the west from Sambirano to 
Onilahy, this fish is known from the following water courses: 
Betsiboka within the region of Maevatanana, lakes of the 
central-west coastal region (Sahapy, Tsianaloka, Bemamba), 
the Mahajanga region and St Augustin near Toliara. Known 
also from the Pangalanes. On the east coast, it is much rarer 
and is usually caught singly. A. madagascariensis 

apparently mounts water courses but is often stopped by the 
first waterfalls. Biology : Anadromous, entering freshwater 
to breed, although a period in salt water is necessary for 
maturation. Migrations are not as regular as some other 
species and take place in large clean western rivers. Mature 
fish measure 25 -30cm. A mouth brooder which only breeds 
once a year, producing 45-80 spherical eggs from October to 
the end of November. Fishing takes place using fixed and 
movable traps, nets and occasionally by line. Very 
important fishery, catches may be smoked or sold fresh but 
are rarely salted. Markets in Antananarivo are an important 
outlet for smoked fish. In 1983, the sale price was about Mg 
F 1000/kg. The fishery is decreasing. 

Family CYPRINODONTIDAE 
* Pachypanchax homolonotus (Dumeril, 1861). 
Found in small streams and the 'matsabory' of the central 
west and north-west of the island. Considered rare, it is 
localized in the west from Antseranana and Nosy Be to 
Morondava. Records exist from Lac Kinkony, River 
Maroparosy by Mevatanano, a brook at Andrafiavilo, 
Manitrano, freshwaters at Ankarana, brook at Antikotozo, 
marsh at Mihilaka, marsh at Ankirihitra, Maevatanana. 
Aquarium species (11 ) 

* Pantanodon madagascariensis (Arnoult, 1963). 
Formerly Orvzias . Found in a few forested hill streams 
around Mahambo, Tampolo-Fenoarivo Atsinanana, and the 
east coast in acid water of pH 6. It is likely to be more 
widely distributed than this in the eastern coastal forest 
region without being anywhere abundant. Considered rare. 

Family ATHERINIDAE 
* Bedotia geayi Pellegrin, 1907. 

Has a wide distribution from the north to the south of the 
island on the eastern side between the coast and 600 m 
altitude. Specific records include: Fort Carnot (River 
Sandranata) Befotaka, (600 m altitude), Karianga (500 m 
altitude), Toamasina, Mahambo, Taolanaro, River 
Mananana (100 m altitude) and the mountain streams of 
Mananjory. It is thought to prefer acid waters. An 
attractive and sought after aquarium species, which is also 
eaten locally. Possibly conspecific with B. tricolor . 



* Bedotia longianalis Pellegrin, 1914. 

Found in similar regions to B. geayi , although preferring 
higher altitudes, ca 450-750 m; recorded from Mahambo to 
Fenoarivo Atsinanana. It generally prefers fresher water 
than B. geayi . Fished in freshwater and brackish water. 

' Bedotia madagascariensis Regan, 1903. 

Freshwater but no type locality known. Found in regions of 
low-mid altitude on the east coast. Records exist for 
Maroansetra and Ambodivoangy. Thought to be rare, this 
fish has a limited distribution. Kiener lists it from the coasts 
of the east-central region, although Arnoult and Bauchot 
found it in abundance in the north-east of the island. 

* Bedotia tricolor Pellegrin, 1932. 

Regions of low-mid altitude on the east coast. Recorded 
from rivulets flowing into the River Faraony (Mahakara 
Province). Attractive species - may be caught for aq larium 
specimens. Possibly conspecific with B. geayi . 

* Rheocles alaotresis (Pellegrin, 1914). 

Found in the shallow basin of Alaotra, high Maningory the 
Ambatondrazaka basin, Mangoro and the region of 
Anjozorobe. It reproduces in the spring, females producing 
100-200 eggs. Formerly fished in the Lake Alaotri. basin, in 
particular the rivers in the north and northwest of the basin 
during the wet season; this fishery has since collapsed. This 
fish was not eaten fresh and is usually dried. 

* Rheocles sikorae (Sauvage, 1891). 

Central Madagascar and mountain streams in eastern 
Madagascar including the forested region of Perinet. Also 
freshwaters around Mangoro. Caught in baskets and 
occasionally by line but it does not play an important 
economic role. 

* Rheocloides Pellegrini Nichols and La Monte, 1931. 
Monotypic genus. Considered rare, it is found in the Andapa 
basin in north-east Madagascar, Lake Alaotra and the 
district of Ambatondrazaka. 

Family AMBASSIDAE 
* Ambassis fontoynonti Pellegrin 1932. 

Relatively uncommon, found in a limited geographical 
location in the central/east and south-east coastal region of 
Madagascar. Known from the region of Manakara, Faraony, 
Rianila and coastal rivulets. 

Family CICHLIDAE 
* Oxylapia polli Kiener & Mauge, 1966. 

Very limited distribution at Marolambo, Toamasina 
province at about 450 m. Strict ecological niche. 
Considered likely to be threatened due to its very restricted 
distribution. An archaic cichlid in a monotypic genus. 

* Paratilapia polleni Bleeker, 1868. 

The most widely distributed native cichlid of Madagascar. It 
exists throughout Madagascar except A) extreme south; B) 
Isalo and plateau of Horombe; C) above 1400-1600 m 
altitude; D) some semi permanent rivers of the southwest 
and certain western zones. There are probably several 
geographical races. Records exist for Toamaisna, Mahanova, 
Imerina, Morafena, Befotako (600 m altitude), Midongy au 
sud (700 m altitude), River Manampetra (500-900 m 
altitude), Karianga (500 m altitude), Antananarivo 
(1000-1100 m altitude), Lake Alaotra, Ankarana, Mantasoa, 
Antsirabe and Rasaobe. Its range has been increased by 
introductions to waters where it would otherwise have been 
extirpated by exceptional cold spells e.g. Lake Itasy, 
although it has since disappeared from the latter. Quite 
common, but not prolific, it is considered to be vulnerable. 
Habitat : The most widespread endemic cichlid and most 
euryhaline, this fish can be found in semi-permanent 
watercourses, streams and rivers, lakes, lagoons and marshes, 
in fresh and brackish waters although it does not approach 
the mouths of rivers. Biology : An adaptable species which 
can cope with large changes in climate and water chemistry. 
In mountainous areas, it is limited by temperature, not 



■193- 



An environmental profile of Madagascar 



Paratilapia polleni (contd.) 
living in waters below 12-13 ' C. Sexually dimorphic, males 
have blue and green/yellow colouring. Slow growers, they 
may reach a maximum of 30 cm and 800 grammes. At the 
first spawning, 800-900 eggs are produced and on 
subsequent spawnings this may rise up to a maximum 
recorded of 3610. Guarded by the parents, incubation is 
controlled by temperature and lasts for 12 days at 22 ' C. 
The nest may be a hollow dug out of the bank. Young fish 
become independent 4-5 weeks after hatching. Parents may 
cannibalize the young. Its diet comprises plankton when 
young, later becoming omnivorous and taking mainly insects 
and aquatic larvae as well as algae. Fisheries : Prized food 
fish which was dominant in fisheries of the past, but now of 
reduced importance as new introduced species compete. 
Notable fisheries were at Alaotra and around Antananarivo. 
Experimental aquaculture has taken place in the paddy 
fields. It is a luxury fish well received in the markets. 
Introduced tilapias have reduced the numbers of this fish. 

* Paretroplus dami Bleeker, 1868. 

Occurs in the north-west at low altitudes. The type 
specimen was collected from Imerina and records exist for 
River Sambirano, Ampombilava, Mahanaro and 
Ambalomainty, Betsiboka, Kamoro, and Mahajamba, Nosy 
Be and Lake Ambanga. In the north, the distribution 
overlaps with P. petiti and in the centre with P. kieneri and 
P. maculatus . Considered vulnerable. Biology : Little 
known, likely to be similar to its congeners. Fisheries : In the 
north-west region this fish does not form an appreciable part 
of the catch. Some smoked fish were exported to 
Antananarivo. 

* Paretroplus kieneri Arnoult, 1960. 

Fairly widespread being abundant in Lake Kinkony and also 
found in the regions of Maevatanana, Ambato-Boeni, 
Tsaramandroso and Kamoro. Overlaps with P. maculatus 
and P. petiti in Kamoro. Less common than P. petiti in 
Lake Kinkony. Considered vulnerable. Habitat : Typically 
freshwater species. In Lake Kinkony it is largely found in 
areas where vegetation is encroaching or where the lake is 
very deep. Biology : Thought to reproduce several times per 
year, the eggs are laid on aquatic vegetation. A tough 
species, it is surviving where P. petiti has disappeared. 
Fisheries : Important fisheries in Lake Kinkony, taken by net 
and occasionally line. Regulated throughout the course of 
the year. Most of the catch is smoked at Mitsinjo, 
Mahajanga and Antananarivo and then will keep up to two 
months. Attempts have been made at aquaculture in the 
west and in warm interior regions, but it is probable that 
clean water, a large space and abundant natural food are 
required. 

* Paretroplus maculatus Kiener & Mauge, 1966. 
The most localised member of this genus, being found in the 
central north-west. Currently known from the region of 
Lake Amparihibe-Sud (where it was abundant in 1966), 
Tsaramdroso, Betsiboka and Kamoro. Overlaps in the north 
of its range with P. dami and with P. kieneri and P. petiti in 
Kamoro. Considered vulnerable. 

* Paretroplus petiti Pellegrin 1933. 

This fish has a wide fragmented range, being known from the 
north-west coastal area down almost as far south as the lies 
Barrren. Type from the River Maintimaso (Mahajanga 
Province). Records from Maintirano, in Ambanja and in the 
interior of the country around Tsamandroso, in particular in 
the small lake of Ampijoroa. It is absent however from 
several intermediate zones eg. Lake Sahapy, Lake Amparihy, 
and Matsabory south of Soalala. In Kamoro, P. petiti may 
previously have been common but is now absent from many 
lakes and rivers. This is probably due to the degradation of 
the water by the lateritic mud carried from the Hauts 
Plateaux as they become increasingly degraded by fire. 
Abundant in Lake Kinkony, neighbouring lakes and in the 
region of Tsaramandroso. Overlaps with P. kieneri and 
P. maculatus in the Kamoro. Considered vulnerable. 
Habitat : Has a preference for thickly vegetated areas and the 
edges of lakes. Typically freshwater. Biology : Grows 
rapidly, reaching 35 cm and exceptionally 40 cm. 



Paretroplus petiti (contd.) 
Reproduces prolifically; a female of 25 cm may produce 2000 
eggs which are pink and are laid in aquatic vegetation. 
Breeding occurs during most of the year and fishermen think 
that the females lay 2-3 times a year. Parents care for the 
young over two to three months. An omnivore, P. petiti 
feeds on plankton, algae, insects and small crustaceans as 
well as the pulp of the tubercles of waterlilies. 

' Paretroplus polyactis Bleeker, 1878. 

Limited to an extensive coastal band from Antseranana to 
Taolanaro and is more common in the canals of the 
Pangalanes. Occupies freshwater except in the rainy season 
when it may enter littoral zones. Type collections were made 
in Toamasina and Imerina. Common in freshwater and 
estuaries. Habitat : Only species of this genus found on the 
east coast where it occurs in rivers, streams, coastal lakes 
and lagoons but rarely enters into the mouths of rivers which 
are too saline. P. polyactis does not range above an altitude 
of 250-300 m. Found in fresh and brackish waters, 
preferring warm, clean, still waters. Biology : Grows rapidly 
and may attain 40 cm in length. Reproducing throughout 
November to March, adults make a hollow depression 
beneath a stump, branch or other object and eggs are laid on 
the underside. The diet is based on plankton, molluscs and 
small shrimps. This probably explains the pink colour of the 
fish flesh. Fisheries : A full bodied good food fish, it is much 
sought after, especially in the markets of Toamasina which 
are supplied by fisheries in Ivoloina, Ivondro, Mahatsara and 
the Pangalanes. In the Pangalanes this fish plays a definite 
economic role. However, the methods of capture never bring 
in large hauls - possibly best as the fish appears to have a 
limited resistance to overfishing. May be raised in ponds 
and many attempts have been made at Ambila-Lemaitso. It 
is likely that this clean water fish, which requires space and 
abundant, natural food, may not be suitable for intensive 
aquaculture. 

* Ptychochromoides betsileanus (Boulenger, 1899). 
Confined to the central and southern central parts of 
Madagascar, this fish is named after the site of its discovery 
in Betsileo. It is also known from Lake Itasy, 

Ambalavao-Fianarantsoa, Mandoto, Haut Matsiatra, 
Ampamaherana, Mananantanana, Zamandao and Ivohibe. 
Abundance : Reportedly in decline. In 1933 it was reported 
that this fish constituted 40% of the species in Lake Itasy 
but has now effectively disappeared as a result of 
competition with introduced tilapine fishes Declines are also 
occurring elsewhere following introductions of tilapia. A 
further introduction, of the water hyacinth, Eichhornia 
crassipes , may also be affecting it by reducing the amount of 
flowing water and preventing reproduction in favoured rocky 
areas. Deforestation in the Hauts Plateaux has changed the 
water regime, reducing the water quality, particularly after 
heavy rain when laterite soil is brought down. It seems 
likely that this fish may have been exterminated in several 
areas and is considered threatened. Habitat : Preferring 
clean oxygenated waters, it may be found in rocky passages 
(around Fianarantsoa) and in waterfalls (near 
Manantanana). This species is mainly found in rivers, rarely 
in lakes. Biology : This fish needs clean, cool, well 
oxygenated water. In rivers it is most abundant in waterfall 
pools up to 2-3m deep and in Lake Itasy it does not occur in 
areas of warm water which are less oxygenated. An 
omnivore P. betsileanus feeds on larval insects, vegetation, 
small fish and shrimps, and is considered to grow more 
rapidly in rivers. There is one spawning period in October 
and the start of November, when females lay up to several 
hundred eggs, by preference on rocks up to 1.5 m diameter 
but, failing that, on sandy shores. Fisheries : One of the best 
freshwater fishes for eating, it has a delicate taste, this fish is 
not suitable for aquaculture due to its exacting 
requirements. Partly due to its slow growth and limited 
reproductive capability, it is rare for fisherman in Lake Itasy 
to capture fish exceeding 300 g or 22 cm in length. A further 
problem is that it is a difficult fish to transport. 



194- 



Appendix 2: fauna! lists 



* Ptychochromis oligacanthus Steindachner 1880. 
There are 4 distinct geographical races. (A = extreme east 
coast, B = north-west and Nosy Be, C = Basin of 
Mandritsara, D = south-west around Tulear). This fish is 
found almost throughout the island and records exist from 
Toamasina, Mahanova, Imerina, Fenoarivo Atsinanana and 
Lake Tongobory, Taolanaro, River Manampatra (500-900 m 
altitude), Ambila Lemaitso, Antikotoza, River Sambirano, 
Mahambo, Tampino, Maroramalona and Lake Rasaobe. 
This fish is the most common of the cichlids after P. polleni , 
although tilapia have greatly reduced its numbers. 
Considered vulnerable. Habitat : Lives in fresh and brackish 
waters, preferring warm or lukewarm water. Although 
tolerant of salty water, it rarely enters river mouths. 
Typical of coastal zones and of the foothills, it is found in 
rivers, streams lakes and lagoons of the east coast. Least 
stenohaline of the cichlids. Prefers large tracts of 
freshwaters which explains its absence from small coastal 
marshes although it is found in intermediate areas. It does 
not penetrate above 300-350 m altitude. An initially 
successful introduction to the small Lake Ambohibao 
(1150 m altitude) failed due to harsh winter conditions. It is 
polymorphic, coastal specimens differing from inland 
specimens by being fatter. Biology : Reaching a maximum of 
27 cm and weight of 500 grammes. A nest of 5-10 cm 
diameter is scooped out of the sandy substrate and 
incubation of the eggs occurs under the supervision of the 
parents. Egg laying continues from November to March and 
egg incubation takes 8-10 days. The alevins stay in shoals 
which may venture outside the nursery area. Omnivorous, 
taking small insects (and also plankton). In the Pangalanes 
shrimps are favoured. Fisheries : In the whole of the east 
coast this fish constitutes an important part of the fisheries 
and may reach 25% of the catch in certain areas. 
P. oligacanthus has been raised in artificial lagoons eg at 
Ambila-Lemaitso. By virtue of its small growth and limited 
resistance, this species is of less value to aquaculture than 
the tilapias. 

Family ELEOTRIDAE 
* Eleotris tohizonae (Steindachner, 1880). 

Found along the east coast and on the northwest coast, 
known locations are Antsirabe, Nosy Lava, source of the 
Amboboko, Ivoloina, Fenoarivo Atsinanana, Mahambo, 
River Tohizona (Foizona), Toamasina, basin of the River 
Mananara, River Faroiny, Mahambo Province and the 
Taolanaro region. It prefers small rivulets at low altitudes. 
Too small to be of economic importance but is nevertheless 
taken. 

* Eleotris vomerodentata Mauge, 1984. 

Known from the Pangalanes to Andevoranto on the east 

coast of Madagascar. 

* Ratsirakia legendrei (Pellegrin, 1919). 

Recently changed genus from Eleotris . Found strictly in 
freshwater at high altitudes (750-1400 m) (11), this fish is 
widely distributed in the Imerina region. It is known from 
Tsaratanana, Mangoro, Lake Alaotra, Perinet, and Midongy 
du Sud although it is rarer in the central south of the island. 
Females lay about 200 eggs, which are guarded by the 
males. Preyed upon by the introduced Micropterus 
salmoide8 . It is too infrequent to be of economic importance. 

* Typhleotris madagascariensis Petit, 1933. 
This blind fish inhabits caves and subterranean waters in the 
southwest of the island. Records exist from Ambilahilalika, 
Malazomanga, Mitoho, Lalio, Andramanaetse, Nikotsy. 
Tsimanampetsotsa, Mitono, Betioky to Soalara, Efoetsy to 
Itampolo. Very limited distribution in the west, therefore 
any disturbances could affect the fish. Considered 

threatened. Eats Crustacea and cave insects. 

" Typhleleotris pauliani Arnoult. 1959. 

Caves and subterranean waters in the southwest of the 
island. Blind. Caves Safara, Andranomaly, Ankilivona, 
Morombe, Baie d'Assassains. Very limited distribution in 
the west, therefore any disturbances could affect the fish. 
Considered as threatened. Feeds on small aquatic insects. 



Family GOBIIDAE 
* Acentrogobius therezieni Kiener, 1963. 

Type locality is a rising spring of fresh calcareous water 
(tsingy) at Antsonjo and is also known particularly from 
limestone formations south of Soalala including the river 
Andranomavbakely (tributary of the Andranamavo) which 
enters the Mozambique channel by Soalala. A scarce fish, 
considered rare. 

Bathygobius samberanoensis (Bleeker, 1867). 
Recorded from Ambavombe (littoral zone) and Bay of 
Bombetake east of Bastard and near Taolanaro. Especially 
in the northwest, it prefers brackish waters. 

Chonophorus macrorhvnchus (Bleeker, 1867). 
Thinly distributed throughout most of the island and 
principally known from the Sambirano region and north of 
the island. Two forms are known, one from coastal regions 
which may mount rivers for considerable distances having 
been found in the Tsibidy near to Kandreho. Possibly 
euryhaline, it is relatively common. It is strictly freshwater 
and is probably adapted to high altitude. It is preyed upon 
by the introduced Micropterus salmoides . 

Gobius polyzona Bleeker, 1867. 
Coastal areas. Sambirano River. 

* Platygobius hypselosoma (Bleeker, 1867). 
Known from the west and north, including the River 
Sambirano and Ivondro. It is infrequently encountered and 
does not play any economic role. 

Platygobius madagascariensis (Bleeker, 1867). 
Occurring throughout most coastal zones, most densely in 
the River Sambirano, this fish occurs in the bay of 
Ampasindava including Nosy Be. It has a secondary 
economic role. 

* Stenogobius verged (Bleeker, 1867). 
Maromandia. Lives in the east coastal region. 

Family KRAEMERIDAE 
Gobitrichinotus arnoulti Kiener, 1964. 

In Madagascar this species is known from the mouths of 
rivers along the central/eastern coast, notably Rianila where 
it is abundant. Appears to inhabit more or less salt free 
sands leaving these to feed when rising water covers them. 
They rebury themselves as the water withdraws. Not of 
economic interest, but known to the fishermen of 
Betsimisaraka. 



B. ENDEMIC SUBSPECIES 

Family ANGUILLIDAE 

* Anguilla nebulosa labiata (Peters 1952) - In Madagascar 
only found in the Lake Itasy region. 

Family CYPRINODONTIDAE 
* Pachypanchax plavfairi (Gunther 1866) - Known from 
Zanzibar and the Seychelles. An attactive species found in 
many localities. Var. sakaramy Holly described as endemic. 
Considered a subspecies of the Seychelles species. Found in 
the extreme north east particularly in the Ambre region. 
Locations include the Foret d'Ambre and Antseranana - 
very localized and needs attention. 

Family ATHERINIDAE 

Hepsetia duodecimalis (Cuvier and Valenciennes, 1837) - 
Known from Ceylon, Malaysia. Var. waterloti Pellegrin, 
1932 - Described from Ruisseau d'Antikotozo, additional 
records are Ambilobe (Antseranana) and the Toamasina 
region. It will extend for many miles upstream and is found 
throughout the Pangalanes Est. It has also been found at 
Ambila-Lemaitso. It is found in mangroves, brackish 
waters, coastal marshes and lagoons. A shoaling species, it 
can only be caught in baskets. 



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An environmental profile of Madagascar 



Family KUHLIIDAE 

Kuhlia rupestris (Lacepede, 1802) - Comores and 
Madagascar. Var. sauvagei Regan considered endemic to 
Madagascar. Found frequently in Onilahy and will enter the 
Sept Lacs and Ambohimatiavelona. Many young have been 
found at Sarodrano (north of St. Augustin) and along the 
west coast. Additional records include Imerina, River 
Ranobe (50km from its mouth), the east coast and the region 
of Toliara. Carnivorous. Reproduction has never been 
observed in freshwater. Their preference for rocky passages 
is utilized by fishermen: traps are set in calm water near 
small waterfalls above which are fixed ant or termite nests. 
The insects fall into the water and attract the fish. Also 
caught in nets. 

Family MUGILIIDAE 
* Agonostomus telfairii Pellegrin, 1932 - Exists on Comoros, 
Reunion, Mauritius and the Seychelles. Var. catalai 
Pellegrin, is considered endemic and is found in a band 
parallel to the east coast including the basin of Ankaibe 
between 100 and 400 m altitude. Also found in Haut 
Mananaire and in rapids on the Mananano. Habitat : This is 
a species which requires strongly oxygenated waters and is 
frequently found in low lying foothill rivers. Typical of clear 
water. Herbivorous. May be caught by line or by net and 
certain fishermen capture large specimens by harpoon. 
Prized flesh and a good sports fish. 



Sicyopterus fasciatus Day 1875 - Found on Mauritius and 
Reunion, it has been considered rare on Madagascar, 
infrequent and abundant in rocky parts of east coast rivers. 
In Madagascar, this fish inhabits rivers in the foothills of the 
east coast usually up to 500 m. Found in strongly 
oxygenated waters often in association with plants such as 
Aponogeton fenestralis and Hydrostachys sp. Migrates to 
the coast. Unusuallly, this fish may use aerial respiration 
and feed on benthic microphages. Fished, but of lesser 
importance than S. lagocephalus . 

Sicyopterus lagocephalus (Pallas. 1770) Found in the 
freshwaters of the Mascarenes. This species has a regular 
migration, entering larger watercourses in large numbers at 
the new moon and for up to 2-3 days afterwards. Found in 
large shoals in coastal waters in the sea and at river mouths. 
Recruitment appears to be variable. Fry are often harvested 
in large numbers and eaten fresh or dried. 

Sicyopterus laticeps (Cuvier & Valenciennes, 1837) - Known 
from River Anjouan (Comoros) and the Mascarenes. 



Family ELEOTRIDAE 
Eleotris ophiocephalus (Cuvier and Valenciennes, 1837) - 
Zanzibar, Mozambique, Comores, Seychelles. In Madagascar 
found in coastal zones including Nosy Be, Nosy Lava. Var. 
madagascarensis is considered endemic and is found in 
coastal waters reproducing in freshwater. 

Family GOBIIDAE 

Gobius criniger (Cuvier and Valenciennes, 1837) - Known 
from E. Africa, India and Malaysia. This is a marine species 
which enters coastal freshwaters. Var. decaryi Pellegrin is 
considered endemic, and has been described from marais 
Fort Dauphin and Mananara. There is no economic interest 
in this subspecies. 

Periophthalmus koereuteri (Pallas, 1770) - Large 

distribution and found from the west coast of Africa to 
Polynesia including Madagascar, Seychelles, India and 
Malaysia. Var. papilio Bloch <fe Sneider 1801 - In 
Madagascar it is found throughout coastal and brackish 
waters especially around rocks and mangroves. An 

amphibious species, it is carnivorous eating insects, small 
Crustacea and fish. 



C. REGIONAL ENDEMICS 

Family KUHLIIDAE 

Kuhlia splendens Regan, 1913 - Suggested endemic but also 
found on Rodrigues and Mauritius. Occasionally found on 
the west side of Madagascar, more rarely on the east side. 
May be fished for although not commonly so. 

Family MUGILIIDAE 

Agonostomus dobuloides (Cuvier and Valenciennes, 1836) 
Considered endemic by, Pellegrin also reports this fish from 
Reunion. Found in the coastal zones of the extreme north 
and north-east of Madagascar. A rarer species found at 
lower altitudes than A. telfairii , some have been captured in 
the river Ankaviabe in the region of Antalahia. Others have 
been found near Antseranana. 



Family BLENNIIDAE 
Sal arias monochrous Bleeker, 



1869 



Listed as endemic. 



Known from Reunion and River Sambirano, Madagascar. 
Not of economic interest. 

Family GOBIIDAE 

Sicyopterus acutipinnis (Guichenot, 1874) - Also found in 
freshwaters in the Mascarenes and in Reunion. 



196- 



MADAGASCAR BUTTERFLIES (except HESPERIDAE) 

Madagascan endemics are indicated by * and Malagasy 
sub-regional endemics are indicated by **. 

Family PAPILIONIDAE 
Graphium evomba r Boisduval, 1836 

'Common over the whole island; not at risk. Endemic to 
Madagascar. 

Graphium cyrnus Boisduval, 1836 

'Common over the whole island; not at risk. Endemic to 

Madagascar. 

Graphium endochus Boisduval, 1836 - RARE 
*An uncommon species which flies on forest edges, often 
feeding on Lantana . Biology unknown but it may breed 
inside the forest. Abundant on Montagne d'Ambre. Its 
status should be monitored. Endemic to Madagascar. 

Atrophaneura (Pharmacophagus) antenor Drury, 1773 

Pharmacophagus is an endemic and monospecific subgenus, 
but the genus Atrophaneura is widespread in the Oriental 
region. A. fP.) antenor is an important endemic, being the 
only Afrotropical representative of the tribe Troidini, the 
Aristolochia-feeding swallowtails of South America (Parides, 
Battus) and the Orient ( Atrophaneura . Troides , 
Ornithoptera etc.). It is probably Madagascar's most 
beautiful butterfly and is sought by collectors. However, it 
is well distributed outside the rain forests and not presently 
at risk. Its status should be carefully monitored. 

Papilio demodocuB Esper, 1798 

An introduced panafrican species, minor pest of citrus. 

Papilio erithonioides Grose-Smith, 1891 

*Mainly in the west and central areas. Not particularly 

uncommon, not threatened. Endemic to Madagascar. 

Papilio grosesmithi Rothschild, 1926 - RARE 

*Mainly in the west. Commercially collected. Requires 

monitoring. Endemic to Madagascar. See data sheet. 

Papilio morondavana Grose-Smith, 1891 - VULNERABLE 
"The rarest of the Malagasy endemics. Threatened by loss of 
habitat and vulnerable to commercial collectors. See data 
sheet. 

Papilio dardanus Brown, 1776 

A panafrican species represented in Madagascar by a special 
race meriones . Not uncommon in the north, east, south and 
south-west. Forest edges. 



Papilio oribazus Boisduval, 1836 
'Quite common and well distributed, 
Endemic to Madagascar. 



except in the west. 



Papilio epiphorbas Boisduval, 1833 

**Well distributed in Madagascar and not at risk; 

the Comoros. 



also on 



Papilio delalandei Godart, 1824 

'Well distributed in forests, especially in the east. Endemic 

to Madagascar. 

Papilio mangoura Hewitson, 1875 - RARE 
An endemic species distributed in the eastern rain forests 
and usually regarded as rare. At present it may not be in 
danger, but deforestation could quickly alter its status. 
Careful monitoring required. Local catchers decoy R 
mangoura with females of the more common P. delalandii . 
which has similar yellow-barred wings. See data sheet. 

Family PIERIDAE 
Catopsiliaflorella F., 1775 
Afrotropical and Oriental. 

Catopsilia thauruma Reakirt, 1866 
"Madagascar and Mauritius. 



Appendix 2: fauna! lists 



Eurema hecabe L.. 1758 
Afrotropical except Cape Province. 

Eurema brigitta Stoll. 1780 

Afrotropical. Ssp. pulchella Boisduval 

Mauritius, Comoro and Aldabra. 



in Madagascar, 



Eurema hapalae Mabille. 1882 

Marshy grassland in tropical Africa and Madagascar. 

Eurema desiardinisi Boisduval, 1833 
•'Madagascar, Comoro Is. 

Eurema floricola Boisduval, 1833 

•'Madagascar, Aldabra, Comoros, Mauritius, Reunion. Ssp 

floricola Boisduval, Madagascar endemic. 

Pinacoptervx eriphia Godart, 1819 

South Africa, Zimbabwe, Botswana, Mozambique, Malawi, 
Tanzania, Madagascar (ssp. mabillei Aurivillius), Sudan, 
Ethiopia, Somali Republic, Kenya, Uganda, Mauritania to 
Senegal, Upper Volta, Niger, Chad, Arabia, Zaire. 

Nepheronia buquetti Boisduval, 1836 

Senegal to Sudan and Ethiopia (ssp. in Niger j, north of 
forest belt, Somali Republic to Mozambique, Zimbabwe and 
Natal (South Africa). Ssp. pauliani Bernardi endemic to 
western and southern dry areas of Madagascar. 

Colotis Calais Cramer, 1775 

Africa south of the Sahara and southern Arabia; absent from 

Cape. Ssp. crowlevi Sharpe in Madagascar. 

Colotis zoe Grandidier, 1867 
'Madagascar endemic. 

Colotis guenei Mabille. 1878 
'Madagascar endemic. 

Colotis evanthe Boisduval, 1836 
"Madagascar and Comoro Is. 

Colotis mananhari Ward, 1870 
'Madagascar endemic. 

Gideona lucasi Grandidier, 1867 
'Madagascar endemic monospecific genus. 

Belenois grandidieri Mabille, 1878 

"Madagascar and Aldabra, replacing B. zochalia . of which 

it could be a race. 

Belenois aurota F., 1793 

Afrotropical and parts of Palaearctic and Oriental. 

Speciemens from Madagascar are strongly marked with 

yellow. 

Belenois creona Cramer, 1776 

Senegal to Nigeria, Sudan, Ethiopia, Somalia, Comoros, 
Arabia, East Africa and Zaire to Cape, Madagascar (endemic 
ssp. prorsus Talbot). 



Belenois helcida Boisduval, 
'Madagascar endemic. 



1833 



Belenois antsianaka Ward, 1870 
'Madagascar endemic. 

Belenois mabella Grose-Smith, 1891 
'Madagascar endemic. 



Dixeia charina Boisduval, 1836 
Kenya to southern Africa, 
endemic to Madagascar. 



Ssp. n arena Grose-Smith 



Appias sabina Felder Si Felder, 1865 

Western Uganda to Nigeria and Sierra Leone, East Africa to 
Malawi and Zimbabwe, Comoros (ssp. comorensis Talbot), 
Madagascar (ssp. confusa Butler). 



•197- 



An environmental profile of Madagascar 



Appias epaphia Cramer, 1779 

Senegal to Zaire, Uganda, eastern and southern Africa, 

Comoros, and Madagascar (endemic ssp. orbona Boisduval). 

Mylothris splendens Le Cerf, 1926 
'Madagascar endemic. 



Mylothris smithii Mabille, 1879 
'Madagascar endemic. 

Mylothris phileris Boisduval, 1833 
'Madagascar endemic. 

Leptosia nupta Butler, 1873 

Nigeria to Angola, Zaire, Uganda, Tanzania, 

endemic ssp. viettei in Madagascar. 



Ethiopia, and 



Leptosia alcesta Stoll, 1784 

West Africa, Ethiopia, Sudan, Uganda, Zaire, Kenya, 
Tanzania to Natal (South Africa). Ssp. svlvicola Boisduval 
endemic to Madagascar. 

Family NYMPHALIDAE: Danainae 
Danaus chrysippus L., 1758 
A common cosmopolitan species. 

Amauris nossima Ward, 1870 - RARE 

*One of the great rarities of Madagascar, distributed mainly 
in the eastern rain forests. Three forms are known. 
Montagne d'Ambre an important locality for disjuncta . 
Believed to be endemic to Madagascar, but D'Abrera lists it 
as possibly occurring in the Comoros. 

Amauris phoedon F., 1798 

* 'Mauritius and possibly Madagascar. Common around the 

coast in Mauritius, but only doubtfully recorded from 

Madagascar. 

Family NYMPHALIDAE: Charaxinae 

Charaxes is a genus of large tailed butterflies, much sought by 
collectors. Almost exclusively forest -dwellers, their biology is 
very poorly known. Adults seek ripe fruits and excrement and 
are trapped by baiting. Their distribution in Madagascar is 
poorly known. Their status needs to be carefully monitored as 
the felling of rain forest proceeds. 

Charaxes cacuthis Hewitson, 1863 

*A subspecies in Paulian (1956) but given full species rank in 
D'Abrera (1980). Well distributed except in the north. 
Endemic to Madagascar. 

Charaxes andara Ward, 1873 

'Well distributed in woodlands and montane areas, but 

apparently not in the forests of the east. Endemic to 

Madagascar. 

Charaxes andranodorus Mabille, 1884 

'Very large (10-12 cm wingspan). Central and northern 
forests, including Montagne d'Ambre. Insufficient data. 
Endemic; restricted in range. 

Charaxes phraortes Doubleday, 1847 

*A forest species. No information available except 

descriptions. Possibly rare. Endemic to Madagascar. 

Charaxes analava Ward, 1872 

'Well distributed except in the south. Apparently a forest 

species. Little information. Endemic to Madagascar. 



Charaxes antamboulou Lucas, 1872 

'Well distributed except in the north. 

Madagascar. 



Endemic to 



Charaxes cowani Butler, 1878 - RARE 

'Apparently very restricted in range, known only from the 
central region, 35 km south of Amboistra at 1700 m. More 
data required. Endemic to Madagascar. 



Charaxes zoolina Westwood, 1850 

Panafrican with a subspecies, betsimisaraka in Madagascar. 
Not at risk, particularly common in the eastern forest (valley 
of Faraony, etc.), and reaching high altitudes in Sambirano. 

Euxanthe madagascariensis Lucas, 1843 - RARE 
*A forest species, rare and with unknown biology. Currently 
well distributed but requires monitoring as deforestation 
proceeds. Endemic to Madagascar. 

Family NYMPHALIDAE: Nymphalinae 
Smerina manoro Ward, 1871 - RARE 

'Endemic monospecific genus. Found in cliffs above 
Toamasina. More data needed. 

Phalanta phalantha Drury, 1773 

Asia and Afrotropical cosmopolitan. P. 2- aethiopica occurs 

widely on Indian Ocean islands, including Madagascar. 

Phalanta madagascariensis Mabille, 1887 

'Endemic to Madagascar. Widespread except in the south. 



Phalanta eurvtis Doubleday, 1847 

Throughout tropical Africa, south to Natal, 

Madagascar and Comoros. 



including 



Apaturopsis kilusa Grose-Smith, 1891 - RARE 

*A strictly forest species found only in north-western 

Madagascar. Very rare and poorly known. Endemic to 

Madagascar. 

Hypolimnas misippus L., 1764 

A Panafrican, Oriental and Australian species. Common in 
the eastern and central areas, of Madagascar, especially on 
the plateaux. 

Hypolimnas bolina L., 1758 

A Panafrican, Oriental and Australian species, common in 

east and central Madagascar. Also on Socotra and Mauritius. 

Hypolimnas dexithea Hewitson, 1863 

'Endemic, but common and widespread in Madagascar. In 

Montagne d'Ambre and elsewhere it flies on the forest edge. 

Hypolimnas deceptor Trimen, 1873 

Coastal forests of eastern Africa from Natal (South Africa) to 
southern Somali Republic, inland to eastern Zimbabwe and 
Malawi, with a separate ssp., deludena Grose-Smith, from 
central and south-western Madagascar. 

Hypolimnas dubius Palisot de Beauvois, 1806 

Panafrican, Hypolimnas d. drucei flies on Madagascar, the 

Comoros and Mauritius. Common. 

Salamis anteva Ward, 1870 

'Widespread endemic in the forests of Madagascar. 

Salamis angustina Boisduval, 1833 

"The nominate ssp. is from Reunion, with a few doubtful 
records and or occasional vagrants to Madagascar. Ssp. 
vinsoni Le Cerf is very rare and possibly extinct in Mauritius. 

Salamis duprei Vinson, 1863 
'Widespread Madagascar endemic. 

Junonia (=Precisl oenone L., 1758 

Entire Afrotropical region, with ssp. epiclelia Boisduval on 

Madagascar, Aldabra, Astove, Assumption and Cosmoledo Is. 

Junonia hierta F., 1798 ( P. lintengensis in Paulian 1956) 
Oriental and Afrotropical regions, with ssp. paris Trimen 
flying in central, southern and south-eastern Madagascar. 



Junonia eurodoce Westwood, 1850 
'In forest clearings. Widespread 
Endemic to Madagascar. 



except in the west. 



Junonia rhadama Boisduval, 1833 
"Madagascar, Mascarene, Comoro 
Doubtfully recorded from Mozambique. 



and Astove Is. 



198- 



Appendix 2: faunal lists 



Junonia goudoti Boisduval, 1833 
"Madagascar and Comoro Is. Widespread. 

Junonia natalica Felder, 1860 

Cosmopolitan. Recorded once from Madagascar. 

Junonia andremiaja Boisduval,. 1833 

*A forest species widespread except in the west. Endemic to 

Madagascar. 

Junonia fPrecisl orythyia L., 1758 
Cosmopolitan. Widespread. 

Vanessa cardui L., 1758 
Cosmopolitan and common. 

Antanartia borbonica Oberthiir, 1880 

**Nominate ssp. on Reunion and Toamasina region of 
eastern Madagascar. A separate ssp., mauritiana Manders, 
is found in Mauritius. 

Antanartia hippomene Hiibner 1823 

Cape Province to Natal and Transvaal in South Africa, with 

a separate ssp., madegassorum Aurivillius, confined to 

Madagascar. 

Byblia anvatara Boisduval, 1833 

Panafrican, the nominate Malagasy subspecies also occurs in 

the Comoros and Glorioso Is. 



Neptidopsis fulgurata Boisduval, 1833 

Coastal forest-savanna mosaic in Kenya and Tanzania, 

the nominate ssp. endemic to Madagascar. Widespread. 



with 



Eurytela dryope Cramer, 1775 

Sub-Saharan Africa and south-western Arabia, 

lineata Aurivillius confined to Madagascar. 



with ssp. 



Eurytela narinda Ward, 1872 

*Widespread in Madagascar except in the east. Closely 

related to E. dryope , possibly a race, in which case E.d. 

lineata would become a race of narinda .. Endemic to 

Madagascar. 

Saliva (= Crenis) howensis Staudinger, 1886 
"Endemic to Madagascar. Well distributed. 

Sallya amazoula Mabille, 1880 

*Well distributed. Endemic to Madagascar. 



Sallya madagascariensis Boisduval, 1833 

'Forests of Madagascar, and probably woodlands too. 

distributed. Endemic to Madagascar. 



Well 



Cvrestis camillus F., 1781 

A Panafrican species with a Malagasy endemic subspecies 

elegans . Widely distributed in Madagascar. 

Neptis decaryi Le Cerf, 1928 - RARE 

*Biology of the Malagasy Neptis species unknown. N. 
decaryi is only known from the type specimen from 
Tsantsany. More data required. Endemic to Madagascar. 

Neptis saclava Boisduval, 1833 

Widespread in sub-Saharan Africa. Subspecies saclava well 

distributed and common throughout Madagascar. 

Neptis metella Doubleday & Hewitson, 1850 - RARE (in 
Madagascar only) 

Panafrican species, but endemic subspecies gratilla only 

known from Toamasina in eastern Madagascar. Very rare. 



Neptis kikideli Boisduval, 1833 
*Well distributed except in 
Madagascar. 



the north. Endemic to 



Neptis sextilla Mabille, 1882- INDETERMINATE 

*Only known from the eastern forests of Madagascar and 

apparently very rare. Very little preserved material is 

available. More study required of this rather enigmatic 

species. 

Cvmothoe lambertoni Oberthiir, 1923 

*A forest species from Sambirano, Montagne d'Ambre and 

the south-east. Biology unknown. Endemic to Madagascar. 

Aterica rabena Boisduval, 1833 

*A forest and woodland species with a wide distribution in 

Madagascar. Endemic to Madagascar. 

Pseudacrea lucretia Cramer, 1775 

Panafrican. Subspecies apaturoides Felder is confined to 

Madagascar, where it is widespread in woodlands. 

Pseudacraea glaucina Guenee, 1864 

'Eastern and southern Madagascar. Presumably a woodland 

and forest species. Apparently common. Endemic to 

Madagascar. 

Family NYMPHALIDAE: Acraeinae 

The genus Acraea is essentially restricted to woodlands and 

forests, but does well in bushy gardens. 

Acraea ranavalona Boisduval, 1833 

**Very common and widespread in Madagascar. Also on the 

Comoros and Aldabra group (Astove Island). 

Acraea hova Boisduval, 1833 - RARE 

'Eastern, central and Sambirano districts. Apparently 

always rare. Endemic to Madagascar. 

Acraea dammii Vollenhoven, 1869 
"Widespread on Madagascar and the Comoros. 

Acraea cuva Grose-Smith, 1889 

Coastal districts of Kenya and Tanzania to Mozambique and 

Malawi. The endemic ssp. villetei flies in Madagascar. 

Acraea igati Boisduval, 1833 

"Very common within its range in Madagascar. Also on the 

Comoros. 

Acraea fornax Butler, 1879 

'Widespread, ecept in the north. Endemic to Madagascar. 

Acraea strattipocles Oberthiir, 1893 

'More limited distribution in eastern and central areas. 

Endemic to Madagascar. 

Acraea masamba Ward, 1872 

'Widespread except in the north. Endemic to Madagascar. 

Acraea silia Mabille, 1886 

'Limited distribution in Sambirano and central areas. 

Endemic to Madagascar. 

Acraea sambavae Ward, 1873 - RARE 

'Eastern and central regions of Madagascar. A rare endemic. 

Acraea lia Mabille, 1879 
'Widespread Madagascar endemic. 

Acraea obeira Hewitson, 1863 

Panafrican, with the nominate subspecies endemic to 

Madagascar. 

Acraea zitja Boisduval, 1833 

'Marshy areas throughout Madagascar. Endemic to 

Madagascar. 

Acraea eponina Cramer, 1780 

Common throughout the Afrotropical region, including 

Madagascar and the other islands. 



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An environmental profile of Madagascar 



Acraea encedon L., 1758 

Very widespread throughout the Afrotropical region, 

including Madagascar. 

Acraea turna Mabille, 1877 

*Fairly common and widespread. Endemic to Madagascar. 



Acraea mahela Boisduval, 1833 

** Widespread in Madagascar and the lies Glorieuses. 

be included in A. terpsicore Sharpe, 1902. 



May 



Pardopsis punctatissima Boisduval, 1833 

Very common in moat dry, open habitats throughout the 

Afrotropical region, including Madagascar. 

Family NYMPHALIDAE: Satyrinae 
Gnophodes betsimena Boisduval, 1833 

Endemic ssp. betsimena in Madagascar. Kenya, Ethiopia, 
Uganda, Sudan, Zaire, Angola, Cameroon, West Africa, 
Cape and Natal Provinces in South Africa, Mozambique, 
Zimbabwe, Malawi, Tanzania. 

Of the 56 species recorded by D'Abrera for the genus 
Henotesia , three are endemic to the Comoros, one to Reunion, 
41 to Madagascar. One is found in Mauritius, Reunion and 
Anjouan as well as Madagascar, and ten are restricted to 
eastern and southern Africa. 

Henotesia anganavo Ward, 1871 
'Endemic to Madagascar. 

Henotesia exocellata Mabille, 1879 
*Endemic to Madagascar. 

Henotesia sabas Oberthiir, 1923 
*Endemic to Madagascar. 

Henotesia bicristata Mabille, 1878 
*Endemic to Madagascar. 

Henotesia erebina Oberthiir, 1916 
*Endemic to northern Madagascar. 

Henotesia strigula Mabille, 1877 
'Endemic to Madagascar. 

Henotesia subsimilis Butler, 1879 
'Endemic to Madagascar. 

Henotesia pallida Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia turbata Butler, 1880 (= ornata Oberthiir) 
'Endemic to Madagascar. 

Henotesia parva Butler, 1879 
'Endemic to Madagascar. 

Henotesia angulifascia Butler, 1879 
'Endemic to Madagascar. 

Henotesia avelona Ward, 1870 
'Endemic to Madagascar. 

Henotesia uniformis Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia oxypteron Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia parvidens Mabille, 1879 
'Endemic to Madagascar. 

Henotesia iboina Ward, 1870 
'Endemic to Madagascar. 

Henotesia anceps Oberthiir, 1916 
'Endemic to Madagascar. 



Henotesia turbans Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia cowani Butler, 1880 
'Endemic to Madagascar. 

Henotesia wardiana Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia curvatula Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia antsianakana Oberthiir 1916 
'Endemic to Madagascar. 

Henotesia ankovana Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia ankova Ward, 1870 
'Endemic to Madagascar. 

Henotesia vola Ward, 1870 
'Endemic to Madagascar. 

Henotesia undulans Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia undulata Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia pauper , Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia ankaratra Ward, 1870 
'Endemic to Madagascar. 

Henotesia laetifica Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia grandis Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia laeta Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia houlbertia Oberthiir, 1923 
'Endemic to Madagascar. 

Henotesia narcissus F., 1798 

"Mauritius, Reunion, Madagascar and Anjouan (Comoros). 

Henotesia strato Mabille, 1878 
'Endemic to Madagascar. 

Henotesia fuliginosa Mabille, 1878 
'Endemic to Madagascar. 

Henotesia undulosa Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia obscura Oberthiir, 1916 
'Endemic to Madagascar. 

Henotesia menamena Mabille, 1877 
'Endemic to Madagascar. 

Henotesia maeva Mabille, 1878 

'Endemic to Madagascar. Possibly synonymous with H. 

narcissus fraterna . 

Henotesia aberrans Paulian, 1951 

'Endemic to the Sambirano region of north-western 

Madagascar. 

Henotesia benedicta Paulian, 1951 
'Endemic to central Madagascar. 



-200- 



Appendix 2: fauna! lists 



The genus Houlbertia Oberthiir, 1910 has eight species, all 
endemic to Madagascar. 

Houlbertia pasandava Ward, 1871 

'Endemic to Madagascar. Possibly synonymous with the 

following species. Females unknown to D'Abrera. 

Houlbertia masikoro Mabille, 1877 

'Endemic to Madagascar. Females unknown to D'Abrera. 

Houlbertia andrivola Mabille, 1877 

*Endemic to Madagascar. Females unknown to D"Abrera. 

Houlbertia cingulina Mabille, 1880 
"Endemic to Madagascar. 

Houlbertia perdita Butler, 1878 
*Endemic to Madagascar. 

Houlbertia wardi Mabille. 1877 
•Endemic to Madagascar. 

Houlbertia narova Mabille, 1877 
'Endemic to Madagascar. 

Houlbertia erebennis Oberthiir, 1916 

'Endemic to Madagascar. A distinctive species, bright blue 

on the recto. 

The genus Masoura Hemming, 1964 has five species, all 
endemic to Madagascar. 

Masoura benacus Mabille, 1884 
"Endemic to Madagascar. 

Masoura antahala Ward, 1872 
'Endemic to Madagascar. 

Masoura ankoma Mabille. 1878 
'Endemic to Madagascar. 

Masoura alaokola Oberthiir, 1916 
'Endemic to Madagascar. 

Masoura masoura Hewitson, 1875 
'Endemic to Madagascar. 

The genus Admiratio . Hemming, 1964, is a monospecific genus 
endemic to Madagascar. 

Admiratio paradoxa Mabille. 1879 
"Endemic to Madagascar. 

The genus Heteropsis has two species, both endemic to 
Madagascar. 

Heteropsis drepana Westwood, 1850 
'Endemic to Madagascar. 

Heteropsis antsianakana Oberthiir, 1916 
'Endemic to Madagascar. 

The genus Strabena Mabille, 1877, has 41 species, all endemic 
to Madagascar. 

Strabena smithi Mabille. 1877 
'Endemic to Madagascar. 

Strabena goudoti , Mabille, 1885 
'Endemic to Madagascar. 

Strabena tamatave Boisduval. 1833 
'Endemic to Madagascar. 

Strabena zaniuca Mabille. 1885 
'Endemic to Madagascar. 

Strabena argyrina Mabille, 1878 
'Endemic to Madagascar. 



Strabena sufferti Aurivillius, 1898 
'Endemic to Madagascar. 

Strabena albivittula Mabille. 1879 
'Endemic to Madagascar. 

Strabena excellens Butler, 1884 
'Endemic to Madagascar. 

Strabena albiviltuloides Paulian, 1951 
'Endemic to Madagascar. 

Strabena corvnetes Mabille, 1885 
'Endemic to Madagascar. 

Strabena nive: .ta Butler, 1879 

'Endemic to Madagascar. Treated by Carcasson as a 

synonym of ba^esii . 

Strabena batesii Felder & Felder, 1867 (= nepos Oberthiir) 
'Endemic to Madagascar. 

Strabena parens Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena propinqua Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena affinis Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena frater Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena ibitina Ward, 1973 
'Endemic to Madagascar. 

Strabena vinsoni Guenee, 1872 
'Endemic to Madagascar. 

Strabena consobrina Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena germanus Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena martini Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena dyscola Mabille, 1880 
'Endemic to Madagascar. 

Strabena andriana Mabille. 1885 
'Endemic to Madagascar. 

Strabena rakoto Ward, 1870 
'Endemic to Madagascar. 

Strabena vicina Oberthiir, 1916 

'Endemic to Madagascar. Doubtful species. 

Strabena soror Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena triophthalma Mabille, 1885 
'Endemic to Madagascar. 

Strabena aurivilliusi D'Abrera 1980 
'Endemic to Madagascar. 

Strabena consors Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena mopsus Mabille, 1878 
'Endemic to Madagascar. 

Strabena perroti Oberthiir, 1916 
'Endemic to Madagascar. 



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An environmental profile of Madagascar 



Strabena impar Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena modesta Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena modestissima Oberthiir, 1916 
'Endemic to Madagascar. 

Strabena io Paulian, 1950 
'Endemic to Madagascar. 

Strabena cachani Paulian 1950 
'Endemic to central Madagascar. 

Strabena isaolensis Paulian, 1951 
'Endemic to western Madagascar. 



Strabena andilabe Paulian, 1951 
'Endemic to the Sambirano 
Madagascar. 



region of north-west 



Strabena tsaratananae Paulian, 1951 

'Endemic to the Sambirano region of north-west 

Madagascar. 



region of north-west 



Strabena perrieri Paulian, 1951 
'Endemic to the Sambirano 
Madagascar. 

Strabena mandraka Paulian, 1951 
'Endemic to central Madagascar. 



Family LIBYTHEIDAE 
Libythea labdaca Westwood, 1851 

A widespread species in the western and central forest blocks 
of Africa, known to migrate in large swarms. Ssp. tsiandava 
occurs in north-western Madagascar. 

Libythea ancoata Grose-Smith, 1891 

'Endemic to Madagascar. Carcasson f in litt . to D'Abrera) 
suggests that ancoata may be a race of L. cvniras Trimen, 
1866. However, the latter, a Mauritian endemic, is most 
probably extinct. 



Family RIODINIDAE 
The genus Saribia , 
Madagascar. 



with three species, is endemic to 



Saribia tepahi Boisduval, 1833 
'Endemic to Madagascar. 

Saribia perroti Riley, 1923 

'Endemic to Madagascar. Three ssp. are recognised, perroti 
in central regions, fiana Riley in the south-west and 
ochracea Riley in the north-west. 

Saribia decaryi Le Cerf, 1922 

'Endemic to Madagascar. Flies in forests above 500m. 

Family LYCAENIDAE 
Spalgis tintinga Boisduval, 1833 
'Endemic to Madagascar. 



1898, with two species, is 



The genus Trichiolaus Aurivillius, 
endemic to Madagascar. 

Trichiolaus mermeros Mabille, 1878 
'Endemic to Madagascar. 

Trichiolaus argentarius Butler, 1879 
'Endemic to Madagascar. 



In the genus Hemiolaus Aurivillius, 1923, five of the six species 
are endemic to Madagascar. 

Hemiolaus ceres Hewitson, 1865 
'Endemic to Madagascar. 



Hemiolaus cobaltina Aurivillius 1898 

'Endemic to northern and north-western Madagascar. 

Hemiolaus varnieri Stempffer, 1944 

'Endemic to northern and north-western Madagascar. 

Hemiolaus marvra Mabille, 1887 
'Endemic to Madagascar. 

Hemiolaus margites Mabille, 1899 
'Endemic to Madagascar. 

Hypolvcaena phillipus F., 1793 

Entire Afrotropical region. Ssp. ramonza occurs in 

Madagascar, Aldabra, Cosmoledo and probably the Comoro 

Islands. 

Leptomvrina phidias F., 1793 
'Endemic to Madagascar. 

Virachola antalus Hopffer, 1855 

Very common, especially in open situations, throughout the 
Ethiopian region, including Madagascar and the Comoro 
Islands. The genus is also represented in the Oriental and 
Australian regions. 

Virachola renidens Mabille, 1884 
'Endemic to Madagascar. 

Virachola batikeli Boisduval, 1833 
'Endemic to Madagascar. 

Virachola dinochares Grose-Smith, 1887 

Open woodland and thorn bush in Natal (South Africa) to 
Botswana, Zimbabwe, Mozambique, Malawi, Zambia, 
Tanzania, Kenya and Uganda. Also recorded from northern 
Nigeria and Madagascar. 

Virachola wardii Mabille, 1878 
'Endemic to Madagascar. 

Anthene princeps Butler, 1876 

Deciduous woodland and Acacia scrub throughout most of 
Africa south of the Sahara. Subspecies smithi Mabille is 
endemic to woodlands in Madagascar. 

Cupidopsis iobates Hopffer, 1855 

Nominate ssp. in moist woodlands from Kenya and Uganda 

to Cape Province (South Africa), Angola and Zaire. An 

isolated population occurs in Togo, Benin and Guinea, with 

another in Madagascar. Sssp. uranochroa is found in 

Ethiopia. 

Cupidopsis cissus Godart, 1822 

Moist grassy areas throughout Africa and Madagascar. 

Petrelaea sichela Wallengren, 1857 

Savanna and woodland throughout sub-Saharan Africa, 

with ssp. reticulum Mabille confined to Madagascar. 

The genus Rvsops Eliot, 1973, is a monospecific taxon endemic 
to Madagascar. 

Rvsops scintilla (Mabille, 1877) 
'Endemic to Madagascar. 

Uranothauma artemenes Mabille, 1880 
'Endemic to Madagascar. 

Cacyreus darius Mabille, 1877 

"Endemic to Madagascar and the Comoro Islands. 

Leptotes pirithous L., 1767 

Very common throughout Africa, Madagascar and much of 

Asia and Europe. 

Leptotes rabenafer Mabille, 1877 
'Endemic to Madagascar. 



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Appendix 2: faunal lists 



Zizeeria knysna Trimen, 1862 

Entire continent of Africa, Madagascar and the Seychelles. 

Zizina antanossa Mabille, 1877 

Entire continent of Africa, Madagascar and Reunion. 

Actizera atrigemmata Butler, 1878 
*Endemic to Madagascar. 

Actizera lucida Trimen, 1883 

Open woodland and disturbed habitats throughout southern 
and eastern Africa, as well as Madagascar. Also Cameroon 
and Benin. 

Zizula hylax F., 1775 

Entire continent of Africa, also Madagascar. 

Azanus soalalicus Karsch, 1900 
*Endemic to Madagascar. 

Azanus sitalces Mabille, 1899 

**Nominate ssp. on Madagascar only, ssp. mavotti D'Abrera 

on Mayotte in the Comoros. 

Eicochrysops hippocrates F., 1793 

Shady places near streams throughout Africa and 

Madagascar. 

Eicochrysops pauliani Stempffer, 1950 

""Known only from the type locality, Mt Tsiranana (1500 m), 

Antseranana, northern Madagascar. 

Eicochrysops sanguigutta Mabille, 1879 
* "Madagascar and Grande Comoro only. 

Euchrysops malathana Boisduval, 1833 

Common in open habitats throughout all Africa, Arabia and 

Madagascar. 

Euchrysops osiris Hopffer, 1855 

Open habitats throughout all of Africa, Madagascar and the 

Comoros. 

Euchrysops decarvi Stempffer, 1947 
"Endemic to Madagascar. 

Lepidochrysops turlini Stempffer, 1971 

"Known only from the type locality, Toliara, south-western 

Madagascar. 

Lepidochrysops caerulea Tite, 1961 
"Endemic to Madagascar. 

Lepidochrysops leucon Mabille, 1879 
"Toamasina, Madagascar. 

Lepidochrysops azureus Butler, 1879 

"Toamasina, Antananarivo and Fianarantsoa, Madagascar. 

Lepidochrysops grandis Talbot, 1937 
"Antananarivo and Fianarantsoa, Madagascar. 

Freveria minuscula Aurivillius, 1909 

"Endemic and widely distributed in Madagascar. 



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An environmental profile of Madagascar 

ENDEMIC NONMARINE MOLLUSCS OF MADAGASCAR 

(T) = known only from type locality 
(A) = known also from Comores 

Locality data for widespread species are not necessarily 
complete. 

N.B. The following authorities have been abbreviated: 
Bedoucha (Be); Blanc (B); Fischer-Piette (F.-P.); Garreau de 
Loubresse (G. de L.); Salvat (S); Vukadinovic (V). 

I. TERRESTRIAL MOLLUSCS 

MESOGASTROPODA 
Family HYDROCENIDAE 

Georissa aurata (Odhner, 1919) 

Funereal caves at Catsepe and Tsingy de Namoroka, 

Mahajanga; Cap Diego; stream bank detritus. 

Georissa detrita Bavay Si Germain, 1920 (T) 
Cap Diego. 

Georissa petiti Germain, 1935 
L. Manampetsa. 

Family PUPINIDAE 

Madecataulus goudoti F.-P. Si Be., 1965 
Betsimisaraka; forest. 

Madecataulus (petitp undescribed 
Ianzamaly (Toliara); ravines. 

Family DIPLOMMATINIDAE 

Diplommatina decaryi Bavay Si Germain, 1920 
Single broken specimen. Cap Diego. 

Malarinia hova Haas, 1961 
Chutes de la Mort. 



Family CYCLOPHORIDAE 
Acroptvchia aequivoca (Pfeiffer, 1857) 
Central Si north-east: Ankaratra; 
(Maroantsetra); forest. 



Ambohivoangy 



Acroptvchia bathiei F.-P. & Be., 1965 

Antsingy, Andranamavo, grottes de Salapango (all in 

Ambongo). 

Acroptvchia bigoti F.-P., B. Si S., 1969 
South: Fieherenna, Toliara; mangroves. 

Acroptychia culminans F.-P. Si Be. ,1965 
North: Mt Tsaratanana; 2000m. 

Acroptychia grandidieri F.-P. Si Be., 1965 

South-west and west: St. Augustin (cave); Tsingy de 

Namoroka; Antalaha. 

Acroptvchia metablata (Crosse Si Fischer, 1873) 

North and north-east: Ambanje; reserve de Marojejy; 

Antalaha region; as far south as Antsiranamatso. 2 varieties; 

(700-2000m). 

Acroptvchia milloti F.-P. Si Bedoucha 1965 
Ambongo; Ankarafantsika; Mahajanga. 

Acroptvchia pauliani F.-P. Si Bedoucha 1965 
North: Mt Tsaratanana; 750- 1400m. 

Acroptychia pauper F.-P., B. Si S., 1969 (T) 
North: Mt Tsaratanana. Single specimen. 

Acroptychia pyramidalis Sykes, 1900 
No locality data. 

Acroptvchia tubulare (Morelet, 1861) 
No locality data. 



Boucardicus albocinctus (Smith, 1893) 
Central-east and east: Mahanovo (?Mahonoro); 
Anosibe. 



Perinet; 



Boucardicus angavokelensis F.-P.. B. Si V., 1974 (T) 
Central east: Angavokely. Under dead branches; single 
specimen. 



Boucardicus beananae F.-P. Si Be., 1965 
North-east: Beanana and Maroantsetra 
Antongil; forest. 



the bay of 



Boucardicus boucardii F.-P. Si Be. ,1965 
Single specimen; no locality data. 

Boucardicus milloti F.-P. & Be. ,1965 (T) 

2 specimens only. Ambatofitorahana. 

Boucardicus nanus F.-P. Si Be. ,1965 (T) 

3 specimens only. Ambohivoangy. 

Boucardicus notabilis (Smith, 1892) (T) 
Single specimen. Toamasina. 

Boucardicus petiti F.-P. Si Be. ,1965 

West: Bemahara (near R. Nameroko); Ambongo. 

Chondrocvclus mamillaris (Odhner, 1919) 

West: Katsepe (funereal cave), Mahajanga, l'Ambongo, 

Antsingy, gorges de Salapango; Amparimgidro; cave and 

gorges. 

Cvathopoma diegoensis F.-P., B. and V., 1974 (T) 
North: Cap Diego. 

Cvathopoma pauliani S., 1968 

North: Andamy on Mt Tsaratanana; 750m. 

Cyathopoma waterloti F.-P., B. and V., 1974 (T) 
North: Cap Diego, baie des Amis. 

Cyclotus milloti F.-P. Si Be., 1965 
Single specimen. Ambohivoangy; forest. 

Hainesia arborea (Crosse Si Fischer, 1871) 

R. Tsidsoubou (or Tsiribihina); banks of river. 

Hainesia litturata (Morelet, 1877) 
Single specimen; no locality data. 

Family POMATIASIDAE 
Cyclotopsis milloti F.-P., B. and V., 1974 (T) 
North: cave entry, south of massif de l'Ankara, Mananjeba; 
single specimen. 

Tropidophora alluaudi (Dautzenberg, 1895) 

North: Mt d'Ambre; Cap d'Ambre; Mt des Francais 

(Antseranana). 

Tropidophora alternans (Pfeiffer, 1853) 
Single specimen; no locality data. 

Tropidophora ambilobeensis F.-P., B. Si S., 1969 
Two specimens. North: Ambilobe opposite Nosy Be. 

Tropidophora andrakarakarensis F.-P. Si Testud, 1973 
(T). Single specimen. Forest of Andrakaraka (Antalaha). 

Tropidophora andrapanga F.-P., B. Si S., 1969 

North-east: Andrapangy, Ambanitaza, Marokosa (Antalaha). 

Tropidophora aspera (Potiez Si Michaud, 1838) 

Mainly extreme north: Nosy Be; Antseranana; Cap Diego; 

Orangea Port Leven; Mt des Francais; Tsiribihina. 

Tropidophora balteata (Sowerby,1873) 

South: Taolanaro and Andrahomana (abundant in south); 
Ambongo; Ambila (Toamasina); Sainte-Lucie; Faux Cap; 
Cap Sainte Marie; plateau Mahofaly; Manombo; Betioky; 
Beloha; Tsiombe; Behara. 



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Appendix 2: fauna! lists 



Tropidophora bathiei F.-P.. 1949 
Morondava, Iabohazo south of Mahajanga. 

Tropidophora bemaraensis F.-P., 1949 
Bemaraha (200m); Ambongo. 

Tropidophora besalampiensis F.-P., 1949 

West coast: Antsingy, Besalampy (Cap St Andre); woods. 

Tropidophora betsiloensis (Smith, 1882) 

May not be valid species; Betsileo; woods near L. Alaotra. 

Tropidophora carnicolor (Fulton, 1902) 
South: Andrahomana, Taolanaro. 

Tropidophora castanea (Pfeiffer, 1851) 
No locality data. 

Tropidophora cavernarum F.-P., B. & S., 1969 (T) 
Single specimen. Anjohibe; cave. 

Tropidophora chavani F.-P., 1949 

Gorges de Salapango (Bemaraka); Antsingy. 

Tropidophora chromium (Morelet, 1877) 
No locality data. 

Tropidophora cincinna (Sowerby, 1843) 
No locality data. 

Tropidophora consocia (Pfeiffer, 1852) 

Widespread. Antseranana; Orangea; Toliara; Mananjary; 

Windsor Castle; Mt des Francais; grotte de Cap Diego. 

Trees. 

Tropidophora coquandiana (Petit de la Saussaye, 1852) 
South: Androka and Andrahava (Toliara); Mahofaly coast 
dunes and coastal woods. 

Tropidophora crenulata Fulton, 1902 
South: Taolanaro. 

Tropidophora cuvieriana (Petit de la Saussaye, 1841) 
North: Nosy B6; Ampotsehy; grottes de l'Ankarana; plateau 
de l'Ankara et de l'Anamera; grotte de Simiar (massif de 
l'Ankara), Nosy Mitziou. 

Tropidophora deburghiae (Reeve, 1861) 
East coast: Mananara. 

Tropidophora deliciosa (Sowerby, 1850) 

North: Antseranana; Mt d'Ambre; cirque de Fanitrys 

(Ankarana); Windsor Castle. 

Tropidophora denisi F.-P.. 1949 

South: Antaramaitsy (n.w. of Cap Ste Marie); 

Andrahomana. Coast. 

Tropidophora denselirata F.-P.. B. & S., 1969 
Massif de l'Ankara. 

Tropidophora deshayesiana (Petit de la Saussaye, 1844) 
North: Nosy Be; grottes de l'Ankarana; Ampotsehy; 
Ankara-Analamera; cirque de Fanitrys (Ankarana). Cave 
entries. 

Tropidophora diegoensis F.-P.. 1949 (T) 
Cap Diego. 

Tropidophora dingeoni F.-P., B. & S., 1969 (T) 
Single specimen. Mt Tsaratanana. 

Tropidophora eustola (Crosse & Fischer, 1887) 
No locality data. 

Tropidophora felicis F.-P. & Be. ,1965 
Nosy Be. 

Tropidophora filopura F.-P.. 1949 

Gorges de Salapango (Bemeraha); Ambongo; woods. 



Tropidophora filostriata ( Sowerby. 1873) 
South: Taolanaro. 

Tropidophora fivanonensis F.-P. & Be. ,1965 
Fivanona. 

Tropidophora formosa (Sowerby, 1849) 

East coast: Foulpointe; Fenoarivo Atsinanana; Mananara. 

Tropidophora fulvescens (Sowerby, 1843) 

North: Ramena on the coast (Antseranana); Cap Diego; 
Orangea; Port Leven; Mt des Francais; Ambohibe 
(Ambilobe); Antalaha region. 

Tropidophora fuscula (Pfeiffer, 1851) 

Nosy Be; massif du Manongarivo (Sambirano) (1000m). 

Tropidophora gallorum F.-P., B. & S., 1969 (T) 

Two specimens. Mt des Francais (Antseranana); under trees. 

Tropidophora goudotiana (Sowerby, 1843) 
North east near baie d'Antongil: Beanana, Ambohivoangy, 
Ambohitsitondrona (all in Maroantsetra); Ambodirafia, 
Ambodilalona (both near Antalaha); forest. 

Tropidophora grisea (Pfeiffer, 1853) 
No locality data. 

Tropidophora humberti F.-P., 1949 
Plateaux of Ankara-Analamera; Ankarana. 

Tropidophora interrupta F.-P., B. & S., 1969 

South and north-east: Miary (Toliara); Vohimarina to 

Sambave. 

Tropidophora ivongoensis F.-P., B. & V., 1974 (T) 

East coast: Soanierana-Ivongo, south of bay of Antongil; 

single specimen. 

Tropidophora iohnsoni (Smith, 1882) 
Central north-west: south of Trabonjy. 

Tropidophora lamarcki (Petit de la Saussaye, 1841) 
North: Antseranana; Windsor Castle; Mt des Francais. 

Tropidophora ligatula (Grateloup, 1840) 

North: at least 25 sites from Vohimarina to Antalaha. 

Tropidophora lirata Pfeiffer, 1852 
No locality data. 

Tropidophora microchasma (Pfeiffer, 1856) 
South: Toliara. 

Tropidophora milloti F.-P., 1949 

North and west: Ankarana (cave entry); Nosy Be; Nosy 
Mamoko; isle de Ambaritelo; Antalaha (several localities); 
old walls and ruins; (280m). 

Tropidophora morondavensis F.-P., 1949 (T) 
Single specimen. Morondava. 

Tropidophora moulinsii (Grateloup, 1840) 

North: Nosy Faly, Mananjary, Mt des Francais, 

Antseranana, Sambirano; caves. 

Tropidophora multifasciata (Grateloup, 1840) 

North, north-east and south: Toliara; Andavadoaka; 

Vohimarina; south Sambave; Beraty (Maromandia). 

Tropidophora occlusa (Morch, 1832) 

Extreme north: Ankara-Analamera (forest & cave entries); 

cirque de Fanitrys (Ankara). 

Tropidophora oppessulata F.-P., B. k. S., 1969 (T) 
Single specimen. Antseranana. 

Tropidophora perfecta Fulton, 1903 
South: Taolanaro. 



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An environmental profile of Madagascar 



Tropidophora perinetensis F.-P. & Be. ,1965 

east: Anosibe; Ikongo; PSrinet; Ambohivoangy; forest, 

100-1100m. 

Tropidophora petiti F.-P., 1949 
Gorges de Salapango (Bemaraha). 

Tropidophora philippiana (Pfeiffer, 1852) 
Widespread in south: Faux Cap; Toliara; Antsepoke; Miary; 
Cap ste Marie; L. Manampetsa (dunes); Androka; 
Andrahava; Onilahy; grotte de Lovenobato (on bank of 
Onilahy); St Augustin; Sarodrano; Nosy Katafana; 
Miandrarah (Manombo); banks of Fieherenana; Morombe; 
Mangoky; Nosy Andramona; Beloha; Efoetsa. Coastal dunes. 

Tropidophora principalis (Pfeiffer, 1859) 
No locality data. 

Tropidophora propeconsocia F.-P. & Be. ,1965 (T) 
Single specimen. Mt d'Ambre. 

Tropidophora puerilis F.-P. it Be., 1965 (T) 
Single specimen. Mt Tsaratanana; 1400m. 

Tropidophora pulchella (Sowerby, 1843) 

North-east: Ranolanina (n.w. of Ivontaka); 

Ambohitsitondrona (near bay d'Antongil); 10 localities near 

Antahala; forest, 400-700m. 

Tropidophora pyrostoma (Sowerby, 1843) 

West, central west and north east: Morondava, Bemaraha 

(near Miandrivazo); Antalaha region on coast; Maintirano; 

woods. 

Tropidophora reesi F.-P., 1949 (T) 
Ambongo; calcareous woods. 

Tropidophora reticulata (Adams & Reeve, 1850) 
No locality data. 

Tropidophora salvati F.-P. & Be., 1965 (T) 
Single specimen. Amparimgidro (Mahajanga). 

Tropidophora sarodranensis F.-P., B. & S., 1969 (T) 

3 specimens. Saint-Augustin (Sarodrane); region of 

Antalaha. 



Tropidophora secunda F 
Antsingy. 



-P. & Be., 1965 



Tropidophora semidecussata (Pfeiffer, 1847) 
Many varieties. Widespread: Toliara; Miary; Andavadoaka; 
Bejangoa; Faux Cap; plateau and coast of Mahofaly; plateau 
de Miandraraha; L. Manampetsa (fossil); Onilahy; grotte de 
Lavenambato; Andranovaha (cliff); Morondava; Ampotaka; 
Cap Ste Marie (dunes); ravines d'lanzamaly; Bemaraha; 
Namoroka; Ambongo; Mahajanga; Sambirano; Manambato; 
Efoetsa; gorges de Manambolo; Antsingy; Tsiribihina; 
Betioky; Bas Fiherana; grotte de Sarandrano (St Augustin); 
Ambatofinandrahana; cliffs, calcareous woods. 

Tropidophora semilirata F.-P., B. & S., 1969 
Andranohinaly (Toliara). 

Tropidophora sikorae (Fulton, 1901) 

South and north-west: Taolanaro; Amboaniou (Mahajanga); 

Ananalava caves. 

Tropidophora soul ai an a F.-P. & Testud, 1973 

Area around Antalaha: Andrakata in vanilla plantations; 

Marolambo; Analamaho; Ampampamena. 

Tropidophora surda F.-P., B. ii S., 1969 
Two specimens. Mt d'Ambre; Ankarana. 

Tropidophora tenuis (Sowerby, 1843) 
No locality data. 

Tropidophora thesauri F.-P., 1949 
Orangea (Antseranana). 



Tropidophora tomlini F.-P., 1949 

Massif de 1 'Ankara (south of Menemjeba); cave. 

Tropidophora tricarinata (Muller, 1774) 

At least 29 varieties, see F.-P., 1949, p. 41. Mainly east 
coast: Nosy Borah; Cap Est; Foulpointe; Antsingy; 
Andrapangy; Morovare (Farafanga); Ranolalina; Nosy Be; 
massif de Manongarivo (Sambirano); Nosy Komba; Midongy 
(Farafanga); Mananjary forests of Tintingue and Toamasina; 
Befevona; Ambohitsitondrova (700m) Ambohivoangy; 
Karianga (Farafanga); Mananara; Ivontaka; Fenerive; 
Betampona; Haut Sambirano; Zahamena; Manambato 
(Ambotondozaka); Taolanaro; Orangea; Tanala; massif de 
Mangarivo; foret de Beanana; col d'lvohibe; Ambinanitelo 
(Mahajanga); Tampolo; Mt Tsaratanana; Manantely. 

Tropidophora tulearensis F.-P., 1949 

South: Ravines d'lanzamaly in valley of Fiherenana 

(Toliara). 

Tropidophora vesconis (Morelet, 1860) 
Antseranana; Port-Leven. 

Tropidophora vexillum (Sowerby, 1873) 
No locality data. 

Tropidophora vignali F.-P., 1949 

West and north: Mt d'Ambre; gorges de Salapango 

(Bemaraha); Antsingy; Ambongo; Antalaha region; caves. 

Tropidophora virgata (Sowerby, 1843) 

North: Mt des Francais, Anosiravo (Antseranana) south of 

Sambave. 

Tropidophora virgo (Pfeiffer, 1853) 

North: Mt des Francais, Antsirane (Antseranana); cave. 

Tropidophora vittata (Sowerby, 1843) 

Islands and coast off Port-Leven; Nosy Be. woods and sandy 

areas. 

Tropidophora vittelina (Pfeiffer, 1852) or Sowerby 1843?. 
No locality data. 

Tropidophora vuillemini F.-P., B. & S., 1969 
Mt Tsaratanana; 1600- 1800m. 

Tropidophora winckworthi F.-P. ,1949 

Mt d'Ambre; massif de l'Ankara, south of Manemjeba; caves 

and trees, 1100m. 

Tropidophora zonata (Petit de la Saussaye, 1850) 
Extreme south to Antsingy in west and Didy in east: incl. 
Reserve de Marojejy and Ambodilalona (Antalaha); 
Mangabe; forest of Ankaroaka. 

Family ASSIMINEIDAE 
Acmella parvula (Morelet, 1877) 

Centre and east: Taolanaro; Toamasina; Nosy Be; Anjouan; 
Ananalava; Antsirabe, (peat -bog); Vatomandry; Mananara; 
Mananjary; often coastal. 

' Assiminea' geayi Lamy, 1909 
Toliara coast. 



Omphalotropis arbusculae F.-P 
Ambohivoangy; bushes. 



B. & S., 1969 (T) 



Omphalotropis madagascariensis Germain, 1921 

East & south: Andrahomana, Antseranana, Ambovombe, 

several localities around Antalaha, Taolanaro; forest. 

Omphalotropis ripae F.-P., B. ii S., 1969 (T) 
Sandrangato (Moramanga); bushes near waterfall. 

STYLOMMATOPHORA 
Family VERONICELLIDAE 

Desmocaulis subaspera (Fischer, 1883) 

Nosy B£; Nosy Komba. 



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Appendix 2: faunal lists 



Drepanocaulis plateia (Simroth, 1913) 
Nosy Be. 

Drepanocaulis tetragonalis (Simroth, 1913) 
Nosy Be. 

Imerinia excisa (Simroth, 1913) 
East: Sakana. 

Imerinia fischeri (Dupouy, 1966) 

Mountainous region north of Antananarivo; Andringitra. 

Imerinia geayi (Germain, 1918) 
Fiherenana. 



Family ENIDAE 
Edouardia rufoniger (Reeve, 1849) 
North: Mt d'Ambre; Antankaratra I 
Antseranana; cave at Orangea. 



= Antankara); bay of 



Edouardia vesconis (Morelet, 1860) 

Single specimen. Port-Leven on north point; sandy beach 

under dead leaves. 

Rachis ambongoensis F.-P., 1964 

central west coast: Ambongo, Maintirano, Morondava. 

Rachis nigrilineatus (Reeve, 1849) 
East coast: Betsileo and Toamasina. 



Imerinia grandidieri (Crosse & Fischer, 1871) 

Morondava; Nosy Be; Antokofotsy; vallee du Saint Augustin. 

Imerinia hovarum (Robson, 1914) 
Toamasina; Marodotatia. 

Imerinia laevimarginata (Simroth, 1913) 
North and west: Mahajanga. 

Imerinia madagascariensis (Simroth, 1913) 
East: Alaotra. 

Imerinia margaritifera (Heynemann, 1885) 
Central region. 

Imerinia ochracea (Simroth, 1913) 
South west: Fiananarantsoa. 

Imerinia sulfurea (Heynemann, 1885) 
Nosy Be and central region. 

Imerinia verrucosa (Heynemann, 1885) (A) 
Nosy Be; Mayotte (Comores). 

Laevicaulis ocellata (Odhner, 1919) 
Toamasina. 

Sarasinula densinerva (Simroth, 1913) 
Fenoarivo Atsinanana. 

Semperula lilacina (Simroth, 1913) 
West: Sakana; Ste Marie. 

Family VERTIGINIDAE 
Nesopupa decarvi F.-P. & Be. ,1965 (T)? 
Baie des Amis, Antseranana; bushes. 

Nesopupa minutalis (Morelet, 1881) (A) 

May be widespread: recorded from Antananarivo and Nosy 

Be. Also Comoros. 

Nesopupa waterloti F.-P. & Be. ,1965 (T) 
Bank of Baie des Amis, Antseranana; bushes. 

Nesopupa sp. 

Nosy Komba (see F.-P. et al. 1975). 

Family ORCULIDAE 
Fauxulus milloti F.-P. it. Be., 1965 
North-east: Ambohitsitondrona; forest, 700m. 

Family CHONDRINIDAE 

Gastrocopta seignaciana (Crosse ic Fischer, 1879) (A) 
(includes G. madagascariensis Bavay & Germain, 1920). 
Probably widespread: Nosy Komba and Nosy Be, Cap Diego, 
Imorona, Ambovombe, Morombe, Taolanaro; also Comoros 
(= G. tripunctum ) and Europa I. 

Family VALLONIDAE 

Pupisoma waterloti F.-P., B. & V., 1974 (T) 
Antananarivo. 



Rachis tulearensis F.-P., 1964 

South-west coastal area: Toliara; Fieherenna; ravines 
d'lanzamaly; Andavadoaka (between Morombe and 
Morondava); Mangoky; Miary; banks of Onilahy; plateau of 
Mahofaly; Ampotaka; often calcareous cliffs. 

Family FERRUSSACIIDAE 
Cecilioides mariei (Crosse, 1880) 
North: Nosy Be; Antseranana: in piles of twigs. 

Family SUBULINIDAE 
Opeas decaryi F.-P., B. & V., 1974 
Single specimen. Unknown locality. 

Opeas soulaianus F.-P. & Testud, 1973 

North-east: Ambodirano & other localities around Antalaha; 

Nosy Be; baie des Amis (Antseranana). 

Subulina manampetsaensis F.-P. & Testud 
South-west: Lake Manampetsa (in reserve, wooded dunes); 
grottes de Lavenombato (Toliara); l'Onilahy (between 
Toliara and Manampetsa). 

Family ACHATINIDAE 
Leucotaenius adami F.-P.. 1963 
Ampotaka, near Cap Sainte-Marie; river banks, woods. 

Leucotaenius bathiei F.-P., 1963 
South-west: Androaka; coastal dunes. 

Leucotaenius crassilabris (Gray, 1834) 
South: Onilahy; wood on calcareous rocks. 

Leucotaenius favannii (Lamarck, 1822) 

South mainly, also few records from east and west: Ikongo; 

Mananjary Bemaraha; Betioky; St. Augustin; Cap Ste-Marie 

(fossil); Faux-Cap; Itampolo; Antaramaitsy; Betaimbolo; 

west shore of Lake Manampetsa; Lamvato; Androka; Beloha; 

Ambovombe. 

Leucotaenius heimburgi (Kobelt, 1901) 
Andrahamana only. 

Leucotaenius laevis F.-P.. 1963 
No locality data. 

Leucotaenius procteri (Sowerby, 1894) 

South-west: Onilahy; L. Manampetsa; Toliara; Efoetsa; 
Anakao (Toliara); Miandraraha (Manombo); coastal dunes 
and woods. 

Family STREPTAXIDAE 
Edentulina alluaudi (Dauteenberg, 1894) 

North: Mt d'Ambre (foret des Rousettesj; Antseranana; cave 
entries south of River Manemjeba (south of massif de 
l'Ankara); Cap d'Ambre; Vohimarina; Windsor Castle; Mt 
des Francais. 

Edentulina arenicola (Morelet, 1860) 

North: Windsor Castle; Port Leven (under dead leaves); 

Orangea (cave). 

Edentulina battistini F.-P., B. &. S., 1975 

West: Toliara; Amparimgidro (Mahajanga); Salapango; 

Bemaraha; Antsingy. 



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Edentulina gaillardi F.-P. & Be., 1965 (T)? 

North: forest of Ankarafantsika, on banks of Lake Tsimaloto. 

Edentulina intermedia (Morelet, 1851) (T) 
North: Port Leven. 

Edentulina metula (Crosse, 1881) 

North: Nosy Komba; Mt Tsaratanana; Ambabovaky 

(Antalaha). 

Edentulina minor (Morelet, 1851) 

Extreme north, coastal: Port Leven; Nosy Komba; 

Vohimarina. 

Edentulina montis F.-P., B. & S., 1975 (T) 
Single specimen, north: Mt des Francais. 

Edentulina nitena (Dautzenberg, 1984) 
North: Mt d'Ambre; Antseranana. 

Edentulina stumpfii Kobelt, 1905 

North: Nosy Be; grottes de Salapango (Bemaraha); Antsingy. 

Gulella andreana F.-P., B. & V., 1974 

Ravines of Ianzamaly (Toliara); caves of Lavenombato 

(Toliara); Antananarivo (on old wall); Morondava (woods). 

Gulella bouchardi F.-P., B. & V., 1974 (T) 

Single specimen. North-east: Andasibe (Antalaha). 

Gulella cerea (Dunker, 1848) 

Comoros; Bruggen (1981) gives this species as endemic to 

Madagascar but F.-P. ii V. (1964) doubt the records. 

Gulella gallorum F.-P., B. & S., 1975 
Single specimen. Mt des Francais. 

Gulella miarvi F.-P. & Be., 1965 (T) 
Miary - calcareous plateau. 

Gulella soulaiana F.-P., 1973 (T) 

Single specimen, north-east: Virembina (Antalaha). 

Pseudelma madagaacariensis F.-P., B. & V., 1974 (T) 
Elma. 

Family ACAVIDAE 
Ampelita alluaudi (Dautzenberg, 1894) (T) 
North: Mt d'Ambre. 

Ampelita atropos (Ferussac-Deshayes, 1851) 

North and south: Antseranana; Mt d'Ambre (1100m, foret 

des Rousettes) Toliara; forest. 

Ampelita bathiei F.-P., 1952 

North: Mt Tsaratanana; 1200- 1800m 

Ampelita battistini F.-P. & G. de L., 1965 
Ambarobe, Andrahomana. 

Ampelita bizonalis Odhner, 1919 
Catsepe; Mahajanga; caves. 

Ampelita caduca F.-P., B. and S., 1975 (T) 
North: Mt Tsaratanana. 

Ampelita calida F.-P., B. and S., 1975 (T) 
North-east: Sambava. 



Ampelita calypso (Pfeiffer, 1861) 
Mananjary; Farafangana; Manakara; 

Antseranana. 

Ampelita capuroni F.-P., B. & S., 1975 
Central north-east: Manerinerina, Anosibe. 

Ampelita cerina (Morelet, 1877) 
North-east coast: Foulpointe, Maroantsetra. 



Marovare; 



Ampelita chlorozona (Grateloup, 1839) (T) 
Beloha; forest; suspended by spiders. 

Ampelita clotho (Deshayes-Ferussac, 1851) 
North: Antseranana; Mt des Francais. 

Ampelita consanguinea (Deshayes-Ferussac, 1851) 
North coast: Antseranana. 

Ampelita covani (Smith, 1879) 

Pic d'Hivohibe (2100m); Ankavana (Betsileo); forets de 

Fivavona, Vakoana and Amindramiova, Andringitra. Forest. 

Ampelita culminans F.-P., 1952 

North: Mt Tsaratanana only; 2200-2350m. 

Ampelita decaryi F.-P., 1952 
West: Antsingy. 

Ampelita denisi Dautzenberg, 1928 
Farafangana. 

Ampelita dingeoni F.-P., B. ic S. 1975 
North and south: Toliara, Mt d'Ambre. 

Ampelita duvalii (Petit de la Saussaye, 1844) 
Several localities in south: Toliara; forest of Andranovory 
(Toliara); ravines d'lanzamaly (Fieheranana); bay of St 
August in; foret d' Ambatofitoharana; foret de Mangidy 
(Fianarantsoa); Mahofaly (dunes); Onilahy (calcareous 
cliffs); forest, woods, calcareous cliffs. 

Ampelita fulgurata (Sowerby, 1838) 

North-east: Ambohitsitandrona (700m), Marofinaritra, 
Ambohifamotsy and Ambatovaky in Antalaha; Ranolalina; 
Fenoarivo Atsinanana. 

Ampelita funebris (Morelet, 1877) 
No locality data. 

Ampelita futura F.-P. & G. de L., 1964 

Manogarivo (Sambirano); Mt Tsaratanana; woods, 

700- 1400m. 

Ampelita galactostoma (Pfeiffer, 1849) 
Nosy Be. 

Ampelita gaudens (Mabille, 1884) 

North <Sc south west: Nosy Be; Mt Tsaratanana; reserve de 

Marojejy, Antalaha region; Toliara. 

Ampelita globulus F.-P., B. & V., 1974 (T) 
Reserve de Marojejy (1500-2000m). 

Ampelita grandidieri F.-P., 1952 
No locality data. 

Ampelita granulosa (Deshayes-Ferussac, 1851) 

North: Mt des Francais (Antseranana); in dead leaves at foot 

of rocks. 

Ampelita hova (Angas, 1877) 

Mananara; He des Nattes (Pandanus); Manambato; 
Ranolalina; Fenoarivo Atsinanana; Tampolo; 

Amparafaravolo; west of Lake Alaotra; Andratambe; 
Marovare; forest. 



Ampelita julii F.-P. & G. de L., 1965 
North: Ambanje, Maroantsetra, several 
Antalaha. 



localities around 



Ampelita katsaensis F.-P. &. G. de L., 1965 
Haut-Bemarivo; plateau du Katsa; woods, 700-1000m. 

Ampelita lachesis (Deshayes-Ferussac, 1851) 
Orangea; Mt des Francais; Antseranana. 



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Appendix 2: faunal lists 



Ampelita lamarei (Pfeiffer, 1853) 

West, from south to baie d'Antongil: Marovare; 

Ambohitsitandrona; Beanana; Ambohivoangy; reserve de 

Marojezy (l500-2000m); Mt Tsaratanana, many localities 

around Antalaha (one of dominant species) Ambanja. 

Forest. 

Ampelita lamothei (Dautzenberg, 1894) 
Mt d'Ambre. 

Ampelita lancula (Ferussac, 1821) 

Foret de Beanana, Fenoarivo Atsinanana (Tampolo); Nosy 

Borah; Ranolalina; Soanierana-Ivongo. Forest. 

Ampelita lanx (Ferussac, 1822) 

North-east: Betampona reserve; Fenoarivo Atsinanana; 
Mananara; Ranolalina; Ambohivoangy (Maroantsetra); Mt 
Tsaratanana; Nosy Borah; foret de Beanana; Foulpointe. 
Woods and forest. 



Ampelita madagascariensis (Lamarck 1822) 

South and north-east: Taolanaro; Andrahomana; 

Mananjary. Forest. 



Behara; 



Ampelita milloti F.-P., 1952 
Gorges de Salapango (Bemaraha) 



Woods, chalk. 



Ampelita namerokoensis F.-P.. 1952 

Gorges de Salapango (Bemaraha); Tsingy de Namoroka 

(Ambongo). 



Ampelita stragulum (Crosse & Fischer, 1873) 

Forests of Ambohivoangy, Maroantsetra, Beanana. Forest. 

Ampelita stumpfii (Kobelt, 1880) 
Nosy Be (Lokobe). Woods. 

Ampelita suarezensis (Crosse & Fischer, 1877) 
North; Antseranana. 

Ampelita subatropos (Dautzenberg, 1894) 
Mt d'Ambre (foret des Rousettes). Forest. 

Ampelita subfunebris (Mabille, 1886) 
No locality data. 

Ampelita subsepulchralis (Crosse, 1868) 

Several forms with no locality data, but one from Ste Marie. 

Ampelita svlvatica F.-P. ic G. de L., 1965 
North-west: Bevazaha (Ankarafantsika). Forest. 

Ampelita unicolor (Pfeiffer, 1846) 
No locality data. 

Ampelita vesconis (Morelet, 1851) 

Port-Leven. Trees among dunes, under dead leaves. 

Ampelita viridis (Deshayes, 1832) 

St-Marie de Madagascar, Ambatouro, Tsaraka. Dry 

marshes. 



Ampelita omphalodes (Pfeiffer, 1846) 

Widespread: Haut Bemarivo; Farafanga; Mahajanga; Nosy 
Be (Lokobe); Taolanaro; Andrahomana; Amparimgidro; 
Ankarafantsika (100 km south of Mahajanga). Woods, 
600- 1000m. 

Ampelita parva F.-P. ic G. de L., 1965 
North: Mt Tsaratanana; 750-2200m. 



Ampelita pauliani F.-P.. 1952 
North: Mt Tsaratanana (Sambirano). 



Woods, 1700-2000m. 



Ampelita perampla Dautzenberg, 1907 (T) 

North west: reserve de Marojejy (15OO-200Om); Analalava; 

Mt Tsaratanana. 

Ampelita percvana (Smith, 1880) 
Ankafana (Betsileo). 

Ampelita petiti F.-P.. 1952 

Massif de l'Andringitra (up to 2500m) (Fianarantsoa). 

Ampelita pfeifferi F.-P.. 1952 

Orangea (Antseranana); Ankarafantsika. 



Ampelita pilosa F.-P. ic G. 
North: Antsirana. 



de L., 1965 (T) 



Ampelita watersi (Angas, 1877) 
South-east coast: Ekongo. 

Ampelita xvstera (Pfeiffer, 1846) 

From south to Perinet and Antananarivo and 1 'Ambongo, 
also north east: Toamasina; Antananarivo; Mangoro; 
Ambongo; Chutes de la Mort and Anosibe; Taolanaro; 
Matitana; Karianga; Andrapangy in north-east; Antalaha 
region; Mandraka; Mananjary; Zahamena. Woods. 



Ampelita zonata F.-P. & G. de L., 1965 
Masakoamena ic plateau du Katsa 
Woods, 600- 1000m. 



(Haut-Bemarivo) 



Clavator anteclavator Germain, 1913 

South: Mahofaly coast; Ankoba-Andrahomana. Dunes. 



Clavator bathiei F.-P. ic S., 1963 

Central mountains: Massif d'Ankaratra; 

Tsiafajavona. Forest, 1800-2000m. 



Manjakatompo; 



Clavator clavator (Petit de la Saussay, 1844) 

South-west: Andranohinaly; Toliara; St Augustin; Mangoky; 

Fiherana; Onilahy; Benenitra. 



Clavator dingeoni F.-P. 
North: Tsaratanana. 



B. and S., 1975 (T) 



Ampelita robillardi (Angas, 1876) 
South: Taolanaro, Andrahomana. 

Ampelita sepulchralis (Ferussac, 1822) 

Many varieties. East and south west: Lake Zanavorany; 
Ambila Le Maitso; Betampona reserve; Sihanaka; 
Soanierana-Ivongo; south of Mahofaly plateau. 

Ampelita shavi (Smith, 1879) 
South-east: Tanala. 

Ampelita soulaiana F.-P. ic Testud, 1973 
North-east: region of Antalaha. 

Ampelita stephani F.-P. ic Testud, 1973 (T) 

Single specimen. Forest of Ambohimitsinjo (Antalaha). 

Ampelita stilpna (Mabille, 1884) 
No locality data. 



Clavator eximius (Shuttleworth, 1852) 

Central-east: Ekongo; Imerina; Antankaratra; Perinet; 

Analamazaotra forest, 300-800m. 

Clavator grandidieri (Crosse ic Fischer, 1868) 
South, mainly near coast: Cap Ste Marie (fossil); Taolanaro 
Andrahomana; Mangoky; gorges du Manombo; Fiherana 
Manampetsa; Mahofaly coast; Ikonka; Tsiobe 

Behara-Bevia; Mananjary; Beloha. 

Clavator iohnsoni (Smith, 1882) 

Centre: Antananarivo; Anonive; Ambositra. Forest, 1800m. 

Clavator moreleti Deshayes- Ferussac, 1851 
North-west from Tsaratanana to Nosy Komba ic Mt 
d'Ambre (foret des Roussettes); Antankaratra 

(=Ankaratra); Ranolalina region; Nosy Be; Antalaha region; 
Maroantsetra; Sambirano. Forest, 700m. 



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An environmental profile of Madagascar 



Clavator obtusatus (Gmelin, 1790) 

South coast and centre: Antananarivo; Taolanaro; 

Andrahomana; Faux Cap; Cap Ste Marie; Mahofaly plateau; 

Betaimbolo. 

Clavator pauliani F.-P. & S., 1963 
North: Mt Tsaratanana; 1500-2200m. 



Clavator praecox F 
South: Toliara. 



-P. & S., 1963 



Clavator watersi (Angas, 1878) 
No locality data. 

Helicophanta amphibulima (Deshayes-Ferussac, 1851) 
East: Fenoarivo Atsinanana to Maroansetra; Nosy Be; 
Sambirano; reserve de Marojejy (1500-2000m); Chutes de la 
Mort; Bevahara (forest of Ankarafantsika); several localities 
around Antalaha; Perinet; Maroantsetra; Beanana; 
Ranolalina; Mananara. Forest. 

Helicophanta betsiloensis (Angas, 1879) 
Tanala, south east of Betsileo. 



Helicophanta bicingulata (Smith, 1882) 

East coastal area: Mananjary; Ekongo; 

Taolanaro; forest of Ranomafana. 



Marovary; 



Helicophanta echinophora (Deshayes-Ferussac, 1851) 
South: Taolanaro; Andrahomana. 

Helicophanta farafanga (Adams, 1875) 
Farafanga; Ekongo. Forest, sand plains. 

Helicophanta geavi F.-P., 1950 

South-east: Betioky; south of Mahofaly plateau; Toliara; 

Beloha; St Augustin; Ambovombe. Coastal dunes and 

bushes. 

Helicophanta gloriosa (Pfeiffer, 1858) 

South: Toliara; Andrahomana; Faux Cap; Tsiribihina; 
Ambovombe; Beloha Taolanaro; forest of Fivanona 
(Andringitra); Mangidy. Dunes, bushes, crops and forest. 

Helicophanta guesteriana (Crosse, 1868) 
East: Marovare (Farafangana). 

Helicophanta ibaraoensis (Angas, 1879) 

May be widespread. East: Perinet; Maintirano; Betsileo; 

Tanala; Ankarampotsy (950m) (Fianarantsoa). Forest. 

Helicophanta magnifica (Ferussac, 1819) 

Central-east and east: Lake Alaotra; Soanierana-Ivongo, 
Tampolo; Antsiatsiaka (Antalaha); Mandraka; marsh of 
Sifotra; Imerina; banks of Marambato; Betampona; 
Analamazoatra (1000m, forest); Zahamena. 

Helicophanta oviformis (Grateloup, 1839) 
West: Bevahara (forest of Ankarafantsika); Maintirano; 
Ambongo; Ste Marie de Marovoay; Ankara-Analamera; 
Antseranana; Mananjeba; massif de l'Ankara (S. 
Manamjeba); Nosy Be; Haut Sambirano; Mahilaka; massif de 
Monongarivo; Mahajanga; Ankatsepe; Ankarantsika (? = 
Anakarafantsika); Tsingy de Nameroko; Antsingy; gorges de 
Salapango; Tsiribihina; Morondava; Beloha. Forest. 

Helicophanta petiti F.-P., 1950 

South-east; Faux Cap; Mangoky; plateau Miandraraha 
(Manombo); Anakao (Toliara); plateau Mahofaly; valley 
Mandrare (Ifotaka). Coastal dunes. 

Helicophanta socii F.-P., B. & S., 1975 (T) 

Ankarana. Rocky calcareous region, probably cave dweller. 



Helicophanta souverbiana (Fischer, 1860) 
Central-east: Ekongo; Marovary (Farafanga); 
Matitana; Ifandana. Forest. 



Mangoro; 



Helicophanta vesicalis (Lamarck, 1822) 

South & south-west: Ankazoabo; Ifandana; plateau 
Maindraraha (Manombo); ravines d'lanzamaly (Toliara); 
Toliara; Andrahomana; Fort dauphin; Manantantely. 
Woods and ravines. 

Family CHAROPIDAE 
Pilula excavata F.-P., B. & S., 1975 (T) 
Single specimen. Mt Tsaratanana. 

Pilula madecassina F.-P., B. & S., 1975 (T) 
Mt Tsaratanana. 

Family EUCONULIDAE 
Euconulus micra (Morelet, 1882) (A) 
Nosy Be; forest of Ankarafantsika; also Comoros. 

Microcystis bathiei F.-P., B. & S., 1975 

Single specimen. Dunes at Lake Tsimanampetsotsa 

(Mahafaly coast). 

Microcystis madecassina F.-P. & S., 1966 (T) 
North: Mt Tsaratanana; 1600m. 

Microcystis nitelloides F.-P., B. & S., 1975 (T) 
North: Mt Tsaratanana. 

Microcystis tangens F.-P., B. & S., 1975 (T) 
Single specimen. North: Mt Tsaratanana. 

Family HELICARIONIDAE 
Bathia madagascariensis Robson, 1914 (T) 
Nameroko (Ambongo). 

Kaliella ahitsitondronae S., 1966 (T) 

North-east Madagascar: Ahitsitondrona 

(?=Ambohitsitondrona); wood, 700m. 

Ka liella milloti S., 1966 (T) 
Antananarivo; bushes. 

Kaliella soulaiana F.-P., 1973 

Cap Est; forest of Andrakaraka (Antalaha). 

Malagrion paenelimax Tillier, 1979 (T) 

Marojejy Mountains 600m; slug-like, reduced shell. 

Tachyphasis milloti F.-P., B. & S., 1975 (T) 

Single specimen. Forest, massif of PAndringitra, south-east. 

Vitrina madagascariensis Smith, 1882 

Betsileo; Mt Tsaratanana (1400-2000m); forest of 
Ambohitantely; forest of Manjakatompo; Ivohibe (2100). 
High forest. 

Family ARIOPHANTIDAE 
Kalidos aequivocus (Robson, 1914) 
Ambongo, Tsingy de Namoroka. 

Kalidos ambilensis F.-P. & Be., 1966 (T) 

Single specimen. Central east coast: Ambila (Toamasina). 

Kalidos amicus F.-P., B. & S., 1975 (T) 
North: Mt Tsaratanana. 

Kalidos anceyanus F.-P. & S., 1966 (T) 
North: Mt d'Ambre. 

Kalidos andapaensis F.-P., B. & S., 1975 

Widespread in north-east: Andapa; Mt des Francais; 
Perinet; Tsaratanana (1500-2000); Fenoarivo Atsinanana; 
Tampolo; Mt d'Ambre; forest 

Kalidos androkae F.-P. &. S., 1965 
South-west and Toliara; dunes. 

Kalidos anobrachis (Dohrn, 1882) 
South-west but no locality data. 



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Appendix 2: fauna! lists 



Kalidos antsepokensis F.-P., B. & V., 1974 (T) 
South west: Antsepoke (Toliara). Coastal dunes. 

Kalidos balstoni (Angas, 1877) 

Central and south-east; Sandrogato (n. Anosibe); 
Ranomafana; (e. of Fianarantsoa); Ekongo?; Farafangana; 
Mamatantely (Antananarivo); Anosibe; Mandraka (bushes); 
Ivohibe (bushes, col & pic at 2100m). Forest. 

Kalidos basalis (Dohrn, 1882) 
No locality data. 



Kalidos humbloti (Ancey, 1902) 

Widespread from north to south: Mt d'Ambre; Tsaratanana; 
Antankaratra; Fanitrys (Ankarana); Ambohivoangy (bushes, 
50-200m); Tampolo forest (Fenoarivo Atsinanana); coast 
between Toamasina and Nosy Borah; Perinet (under bark); 
Mandraka (under stones); ravines d'lanzamaly (Toliara); 
woods, bushes. 



Kalidos lamvi F.-P. & Be. ,1966 
North: Mt des Francais; Anosiravo; 
Antseranana. 



Sakaramy; Locoube; 



Kalidos bathiei F.-P. & S., 1965 

South and south-west: Morondava; Betioky; Andranohinaly; 

Toliara; Ankazoaba; Befandriana (cap St Vincent). Woods. 

Kalidos bournei Robson. 1914 

Central-west area: Tsingy de Namoroka; other woods and 
forests in Ambongo; gorges de Salapanga; Atsingy; 
Bekopaka; Bemaraha; Betampona; forest. 



Kalidos lapillus F.-P. & Be., 1966 

South-west: Andranovaha; Mangoky; Morombe; 

Miandraraha; Nosy Katafana; Onilahy; Lake 

Tsimanampetsotsa; Mahofaly; Amotaka; cliffs. 

Kalidos mangokyanus F.-P. & S., 1965 

South-west: Antaramaitsy; Reantengy; Andavadoaka; dunes 

and calcareous cliffs. 



Kalidos calculus F.-P. & Be., 1966 (T) 
South-west: Andranovaha; cliffs. 



Kalidos merschardti F.-P., B. and S., 1975 (T) 
North: Mt Tsaratanana. 



Kalidos capuroni F.-P., B. & S., 1975 

Cap Est and several localities around Antalaha. 

Kalidos chastelli (Deshayes-Ferussac, 1851) 

Extreme south-west, coastal: calcareous mountains of Bas 

Fiherana; St Augustin; Andranohinaly; Miary; Fiherana; 

Onilahy; Andranovaha; dunes near Lake Manampetsa; 

Andringy. 



Kalidos microlamvi F.-P., B. and V., 1974 

Cap Diego; Mt des Francais; Amparimgidro (Mahajanga) 

Kalidos milloti F.-P. & S., 1966 (T) 
North: Mt Tsaratanana; 1400 -2000m. 

Kalidos montis F.-P. & Be. ,1966 (T) 
North: Mt Tsaratanana; 1500m. 



Kalidos cleamesi (Smith, 1882) 

South-central: Ambohimitombo, Ankafana (Betsileo); 

Sahana (west Madagascar); Mt Tsaratanana; forest. 

Kalidos dautzenbergianus (Ancey, 1902) 
North: Mt d'Ambre. 

Kalidos decaryi F.-P., B. & S., 1975 (T) 
North: Mt Tsaratanana. 



Kalidos delphini F.-P., 
Taolanaro; forest. 



S. & V., 1974 (T) 



Kalidos ekongensis (Angas, 1877) 

Apparently widespread but only known from a few primary 

forest localities: Ekongo; Kandani; Bongolava (forest, under 

dead wood and bark); Lake Alaotra (primary forest); 

Mahajanga. 

Kalidos eos (Dohrn, 1882) 
Central but no locality data. 

Kalidos eucharis (Deshayes-Ferussac, 1851) 
North-east: Sambirano; woods. 

Kalidos fallax F.-P., B. & S. 1975 (T) 
Single specimen. Mt d'Ambre. 

Kalidos feneriffensis (Adams, 1876) 

North-east: Maroantsetra and Fenoarivo Atsinanana; woods. 

Kalidos fuscoluteus (Grateloup, 1840) 
No locality data. 

Kalidos glessi F.-P., B. & S., 1975 (T) 
North: Mt d'Ambre; single specimen. 

Kalidos hestia (Dohrn, 1882) 

Tampolo (Fenoarivo Atsinanana); Ahitsitondrona; forest. 

Kalidos hildebrandti (Dohrn, 1892) (T) 
Single specimen. South Betsileo. 

Kalidos hova (Odhner, 1919) 

Small area on north-west coast: Catsepe; Amparimgidro. 



Kalidos oleatus (Ancey, 1902) 

North-east; reserve of Marojejy (700-2000m); Mananara 
forest (1800m); Mt Tsaratanana; several localities around 
Antalaha; Antsianaka; Zahamena. 

Kalidos oxyacme (Ancey, 1908) 
Antankaratra. 

Kalidos p'peratus (Fulton, 1901) 

Extreme south-east: Taolanaro; Tranomaro (n-e of Androy); 

Amboisarabe; dunes, forest. 

Kalidos profugus (Ancey, 1902) 

Widespread but few specimens have been found: 

Antankaratra; Farafanga; coast between Toamasina & Ste 

Marie. 

Kalidos prominens F.-P. & S., 1966 
Single specimen; no locality data. 

Kalidos propeanobrachis F.-P. & Be. ,1966 (T) 

Massif du Manongarivo (Sambirano)(1000m); Maheva 

(Farahalana). 

Kalidos rufescens (Grateloup, 1840) 
Forest; no locality data. 

Kalidos secans F.-P., B. & S., 1975 (T) 
Single specimen. Mt Tsaratanana. 

Kalidos soulaiana F.-P., 1973 
No locality data. 

Kalidos tenebricus F.-P., B. ii S., 1975 (T) 
North: Mt d'Ambre. 

Kalidos thalia (Dohrn, 1882) 
No locality data. 

Kalidos tranomarensis F.-P., B. & V., 1974 (T) 
Tranomaro, north-east of Androy. 

Kalidos tsaratananensis F.-P. & S., 1966 
North: Mt Tsaratanana; 1500 - 1800m. 

Kalidos tsialangiensis F.-P., B. & V., 1974 (T) 
South west: Tsialangy. 



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An environmental profile of Madagascar 



Kalidos tulearensis F.-P. & S., 1966 (T) 

Single specimen; south: Ianzamaly, Toliara; ravines. 

Macrochlamvs stumpfii Boettger, 1889 

Widespread, but mainly in north; forest of Manjakatompo; 
reserve de Marojejy (700-2000m); several localities around 
Antalaha e.g. forest of Andrakaratra; Antseranana; cave at 
Orangea; Mt des Francais; Ankara-Analamera; massif de 
l'Ankara; cirque de Fanitrys; Nosy Be; Mt Tsaratanana 
(2500m); Bas Sambirana; Mahilaka; Ambohivoangy 
(bushes); Amparimgidro; Toamasina (under stones); station 
Ivohina; banks of Ivondro; Betampona; Bongolava (in dead 
wood); Zahamena; Andringitra; forest; a pest species. 

Sitala acuta F.-P. & S., 1966 

North: Mt Tsaratanana only; 1400- 1800m. 

Sitala amabilis F.-P. & S., 1966 

Central-east: Manjakatompo & Mandraka; bushes. 

Sitala ambovombeensis F.-P., B. & V., 1974 (T) 
South of Ambovombe: detritus on dunes. 

Sitala ankasakasensis F.-P., B. & V., 1974 (T) 
Single specimen: Ankasakasa (Ambongo). 

Sitala antsingiana F.-P., B. & S., 1966 (T) 
Antsingy. 

Sitala brancsiki Boettger, 1892 (T) 
Nosy Be (Lokobe). 

Sitala culminis (F.-P. & S. in F-P, B, & S 1966) 
North: Mt Tsaratanana only; 1500-2200m. 



Family Ampullariidae 
Lanistes ( olivaceus ) grassetti Morelet, 1863 
Mahajanga; freshwater. Common in lower Mangoky region 
and found elsewhere in Madagascar. May be a form of L^ 



Sitala delaportei F.-P., B 
North: Mt Tsaratanana. 



& S., 1975 (T) 



Sitala elevata ( F.-P. & S. in F-P, B, &. S, 1966) 
Mt Tsaratanana (2000m); Sambirano; Ambanja. 

Sitala filomarginata Boettger, 1892 (T) 
Lokobe (Nosy Be). 

Sitala gaudens F.-P. & S., 1966 
Central-east: Manjakatompo; forest. 

Sitala roedereri F.-P., B. & S., 1975 (T) 
North: Mt Tsaratanana. 

Sitala soulaiana F.-P. & Testud, 1973 
Ansahantangata; Amboafotsy; Ambodirano. 

Family UROCYCLIDAE 
Elisolimax bella (Heynemann, 1882) (A) 
Nosy Be; Nosy Komba; Mayotte (Comoros). 

Elisolimax madagascariensis (Poirier, 1887) 

North east: Toamasina; Ivolohina; Fenoarivo Atsinanana 

Antasibe; banana trees. 

Trochonanina milloti F.-P., B. & V., 1974 (T) 
Ahitsitondrona, north of bay of Antongil. 

Trochonanina tulearensis F.-P., B. & V., 1974 (T) 
Single specimen. Ravines of Ianzamaly (Toliara). 



II FRESHWATER MOLLUSCS 

ARCHAEOGASTROPODA 
Family Neritidae 

Clithon spiniperda (Morelet, 1860) 

Antseranana, Nosy Be. 

MESOGASTROPODA 

Family Truncatellidae 
Truncatella guerini Villa, 1841 
Nosy Be; baie des Amis, Antseranana. 



Pila cecillei (Philippi, 1848) 
Antseranana, Mahajanga, 
Toamasina; freshwater. 



Antananarivo, Fianarantsoa, 



Family Thiaridae 
Cleopatra colbeaui (Craven, 1880) 

Antseranana, Toamasina, Nosy Be. Found most frequently 
in central and northern regions. Freshwater; small streams 
in forest. 

Cleopatra grandidieri (Crosse &. Fischer, 1872 
Eastern central Madagascar and isolated localities on west 
coast; Toamasina, Fianarantsoa, Mahajanga; freshwater, 
forest stream poor in dissolved salts. 

Cleopatra madagascariensis Crosse & Fischer, 1872 
Antseranana, Mahajanga, Toamasina; freshwater. 

Melanatria fluminea (Gmelin, 1767) 

Throughout Madagascar, including Antseranana, 

Toamasina, Antananarivo, Fianarantsoa, Toliara, Mangoky 
area, L. Alaotra; freshwater, small streams and lakes. 

Melanatria madagascariensis (Grateloup, 1841) 
Antseranana, Toamasina; freshwater. 

Family Potamididae 
Cerithidea decollata (Bruguiere, 1838) ? 
Antseranana, Toamasina; freshwater. 



BASOMMATOPHORA 
Family Lymnaeidae 

Radix hovarum (Tristram, 1863) ? 

Widespread; freshwater. 

Family Planorbidae 
Afrogyrus starmuhlneri Brown 
Freshwater (Brown, 1978). 

Afrogyrus (apertus) (Martens, 1896) 
Antananarivo; freshwater. 

Anisus crassilabrum (Morelet, 1860) 

crassilabrum ? 

Widespread. Also Comores; freshwater. 



Afrogyrus 



Bulinus bavayi (Dautzenberg) 

Widespread. With B. mariei , is included as synonym of B. 

forskali (Wright, 1971). Also on Aldabra. 

Bulinus liratus (Tristram, 1863) 

Widespread, common in central and south-eastern areas but j 
some records may refer to B. obtusispira . Freshwater; | 
irrigation channels but comparatively rare in rice fields. 

Bulinus mariei (Crosse, 1879) = B. forskali ? E ? 
Widespread; freshwater. Taxonomic status not clear and I 
may be same as B. forskali from Africa. If a separate species, I 
it may supplant B. bavayi eventually (Wright, 1971). 

Bulinus obtusispira (Smith) 

Extensive range in western region including lower Mangoky I 
district, Mahajanga, Basybasy (in s.w.), Tanararive. I 
Common in rice fields and capable of aestivation for at least I 
seven months; rarely found with B. liratus but the two | 
species are easily confused. 



Family Ancylidae 
Ferissia modesta (Crosse, 1880) ? 
Antseranana, Antananarivo; freshwater. 



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NONMARINE CRUSTACEA OF MADAGASCAR 

ANOSTRACA and CONCHOSTRACA 

Several species have been found, but from the information 
currently available it is not possible to draw biogeographic 
conclusions (Paulian, 1961). 

CLAD O CERA 

About 20 taxa are listed for Madagascar, but it is emphasised 
that these undoubtedly represent at maximum a third of the 
total species present (D.G. Frey, unpublished report). None 
are known to be definitely endemic, but studies on other 
members of this group are revealing that many species, while 
not restricted to particular lakes, are restricted to relatively 
small geographical regions. 

OSTRACODA 

Endemic species include (Decary, 1950): 
Cypris decarvi Gauthier, 1933 

Cyprinotus imus Gauthier, 1934 



CALANOIDA 

5 endemic species in 2 genera; one subgenus is endemic and 
includes a cave variety (Paulian 1961). 

Family PSEUDODIAPTOMIDAE 
Pseudodiaptomus pauliani Brehm 1951. 

Canal des Pangalanes, Mananjary. Found only once in 
stagnant water and dense vegetation of Eichhornia . 

Pseudodiaptomus batillipes Brehm 1954 
Taolanaro. Found once with cladocerans. 

Family DIAPTOMIDAE 

Tropodiaptomus (Anadiaptomus^ madagascariensis (Rylov, 
1922) 

Taolanaro, Antananarivo. Considered characteristic of 
plankton of high plateau lakes: Lake Mandroseza, Lake 
Andrianotapahina. 

Tropodiaptomus (A.) madagascariensis poseidon Brehm 1952 
Rivers and lakes north of Mananjeba and near Ambilobe. 



CYCLOPOIDA 

18 endemic species in 9 genera, 3 of which are endemic 
(Dussart, 1982). 

Family CYCLOPINIDAE 
Allocvclopina madagassica Kiefer 1954 
In sand on beach, lagoon at Maroantsetra. 

Family CYCLOPIDAE 
Halicvlops pusillus Kiefer 1954 
In sand in lagoon of Maroantsetra. 

Halicvlops denticulatus Kiefer 1960 
Found once. Brackish water, Manambato. 

Afrocvclops pauliani Lindberg 1951 
Found once. Antananarivo, Besarety . 

Bryocvclops fRvbocyclopsl pauliani Lindberg 1954 

Cave of Andranoboka near Mahajanga. Subterranean in 

stagnant water. 

Bryocvclops (B.) mandrakanus Kiefer 1955 

Bryocvclops ankaratranus Kiefer 1955 

Found once. Waterfall, in moss. Forest of Ambahona in 

massif d'Ankaratra. 



Appendix 2: faunal lists 

Bryocvclops (Haplocvclopsl gudrunae Kiefer 1952 

Found once. Interstitial water in alluvial sand on bank of 

Menarandra at Tranoroa. 

Bryocvclops fH.^ neuter Kiefer 1955 

Banks of lagoon at Maroantsetra, east Madagascar. 

Bryocvclops (H.) correctus Kiefer 1960 

Occurs in interstitial fauna of phreatic water of Sisaony and 

Faratsiho rivers. 

Cochlacvclops ateles Kiefer 1955 

Known only from type locality. Interstitial water of rich 

alluvium in small mountain waterfall at Faratsiho. 

Goniocvclops primus Kiefer 1955 

Wet moss on edge of Cascade du Foly, massif de 

l'Andohahela, 1500m, Taolanaro. Known only from type 

locality. 

Goniocvclops alter Kiefer 1955 

Known only from type locality. Wet moss, Cascade 
d'Ankaramena, on road between Ambalavao and 
Ankaramena at km 506. 

Psammocvlops excellens Kiefer 1955 

Known only from type locality, on road from Ihosy to 

Betroka at km 300 in tributary of upper Onilahy. 

Mesocvclops annae Kiefer 1930 
Antananarivo. 

Mesocvclops insulensis Dussart 1982 
Lake Bemapazo at Nosy Be. 

Mesocvclops pilosus (Kiefer 1930). 

Edge of shallow lakes with rich vegetation; Lake 
d'Andrianotapahina near Ivato, Antananarivo; grotte 
Andranomaly near Mozombi-Andalambazo. Known only 
from Madagascar. 

Thermocyclops neglectus f. major Dussart 1982. 
Several localities; known only from Madagascar. 

Thermocyclopa consimilis pusillus Dussart 1982. 

Pare de Tsimbazaza at Antananarivo; known only from 

Madagascar. 



HARPACTICOIDEA 

Family CANTHOCAMPTIDAE 

A very interesting, if not particularly diverse group, found 

mainly in streams on rocks and in humid moss (Paulian 1961). 

Echinocamptus pauliani Chappuis 1956 

Known only from type locality; moss, massif de 

l'Andohahela, Taolanaro. 

Attheyella (Mrazekiella) meridionalis Dussart 1982. 
Taxonomic status uncertain. Known from type locality only; 
edge of Lac de Mantasoa. 

Elaphoidella aberrans Chappuis 1954 

Moss, forest of Isaka, Taolanaro and waterfall at Mandraka. 

Family PARASTENOCARIDAE 
Parastenocaris variolata Chappuis 1952 
Phreatic water of Menarandra river at Tranoroa. 

Parastenocaris pauliani Chappuis 1952 
Phreatic water of Menarandra river at Tranoroa. 

Parastenocaris forficulata Chappuis 1952 

Sand (psammic), edge of lagoon of Maroantsetra at 

Ambodivoangy. 

Parastenocaris madagascariensis Chappuis 1952 

Sand, edge of lagoon of Maroantsetra at Ambodivoangy. 



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An environmental profile of Madagascar 



Parastenocaris macaco Chappuis 1952 

Sand, edge of lagoon of Maroantsetra at Ambodivoangy. 

Parastenocaris trisaetosa Chappuis 1954 
Lagoon, Lanirano near Taolanaro. 

Parastenocaris arenicola Chappuis 1954 

Sand, rivers Sisaony and Zazafotsy, on road to Ihosy. 

Parastenocaris pusillus Chappuis 1954 
Sand, lagoon, Lanirano near Taolanaro. 



Armadillo fenerivei Barnard 1958 
Fenoarivo Atsinanana. 

Armadillo euthele Barnard 1958 
Fenoarivo Atsinanana. 

Armadillo silvivagans Barnard 1958 
Forest of Tsaramandroso, Ankarafantsika. 

Armadillo ankaratrae Barnard, 1958 
Forest of Manjakatompo. 



Parastenocaris gracilis Chappuis 1954 
Sand, lagoon of Maroantsetra. 



BATHYNELLACEA 



Bethalus carinatus Budde-Lund 1904 
Antananarivo; Manjakatompo; Antanimeno. 

Bethalus bipunctatus Barnard 1958 
Perinet. 



3 species of syncarids (Parabathynellidae) known from littoral 
phreatic water (Paulian, 1961). 



ISOPODA 



Akermania svlvatica Barnard 1958 
Manjakatompo. 

Akermania hystrix Barnard 1958 
Perinet. 



The main group of terrestrial crustaceans, the wood lice. 52 
species were listed by Barnard (1958) who considered that 
there are still probably many more to be discovered. The 
Ankaratra mountains are especially rich for Isopoda but this is 
probably largely because this area has been most intensively 
studied. A large volume of isopod material is still awaiting 
study (Paulian, 1983). There are several cave species. 

Family STYLONISCIDAE 
Styloniscus albidus Vandel 1952 
Forest of Manjakatompo, Ankaratra Mountains, 2000m. 

Styloniscus vandeli Barnard 1958 
Ambatolaona and Ambanja, 75m. 

Family CIROLANIDAE 

Anopsilana poissoni Paulian &. Delamare Deboutteville, 1956 
Subterranean. Cave of Mitoho, south of Toliara. Blind and 
unpigmented, may be parasitic on blind fish. 

Family TRICHONISCIDAE (ONISCIDAE) 
Madoniscus termites Paulian de Felice, 1950 
East: forest of Tampolo. May be an endemic genus; found in 
termite galleries of Eutermes nigrita , associated with 
Captotermes truncatus . 

Suarezia heterodoxa Dollfus 1895 
Fenoarivo Atsinanana; Mount d'Ambre. 

Suarezia differens Barnard, 1958 

Forest of Manjakatompo (Ankaratra Mountains, 2000m); 

Perinet. 

Didima humilis Budde-Lund, 1909 
Antananarivo; forest of Manjakatompo; Pe'rinet. 

Philoscia reducta Barnard 1958 
Perinet. 

Tura testacea Budde-lund 1902 
Mahajanga; Aldabra. 

Ankaratridium caecum Paulian de Felice, 1950 

Massif de 1'Ankaratra; found under stones near pisciculture 

station at Manjakatompo. Blind; endemic genus. 

Microcercus rotundifrons Barnard 1958 
Forest of Manjakatompo. 

Microcercus mascarenicus Barnard 1958 
Forest of Manjakatompo. 

Armadillo otion Barnard 1958 
Perinet. 



Calmanesia erinaceus Barnard 1958 
Perinet; forest of Niagarakely (Anosibe). 

Calmanesia lonchotes Barnard, 1960 

East central; Moramanga district on edge of forest road. 



AMPHIPODA 

At least 6 endemics (Paulian, 1961). 

Family GAMMARIDAE 
Austroniphargus bryophilus Monod 1925 

Pic Boby in the massif of Andringitra, confined to surface 
layers of water in pools of thick moss among granitic rocks 
(Paulian, 1961; Griveaud, 1981); small, blind. 

Austroniphargus starmuehlneri Ruffo 
Springs near Taolanaro; crenophilic. 

Dussartiella madegassa Ruffo 1979 
Spring water, central Madagascar. 

Family AORIDAE 
Grandidierella mahafalensis Coutiere 1904 
Moheli; Lake Tsimanampetsotsa. 

Family TALITRIDAE 
Orchestia ancheidos Barnard 1916 

Lake Tsimanampetsotsa on banks and in pools with | 
vegetation; Itampolo, a small, brackish lagoon separated 
from the sea by a dune, with dense vegetation. 

Family MELITIDAE 
Melita nitidula Ruffo 1958 
Soalava (south). 

Family ISAEIDAE 
Photis distinguenda Ruffo 1955 
Anove River, north of Toamasina; variable salinity. 



DECAPODA 



Family ATYIDAE 

20 species in genus Caridina , most of which are endemic; 21 
troglobytic genera, Typhlopatsa and Parisia (Paulian, 1961)1 
which include several unique species. Some of the species in I 
the genus Caridina are used for food. 

Caridina angulata Bouvier 1905 

Ranofotsy River, near Fianarantsoa and Lake Itasy. 

Caridina calmani Bouvier 1919 

Ambatonharanana, near Lake Alaotra, (restricted range). 



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Caridina edulis Bouvier 1904 

Malagasy caridina, Saltarelle malgache. Large numbers sold 

cooked in markets. 

Caridina hova Nobili 1905 
Taolanaro. 

Caridina isaloensis Coutiere 1899 
Several localities. 

Caridina lamiana Holthuis 1965 
North-east and east. 

Caridina madagascariensis Bouvier (probably synonym of 
Caridina isaloensis ) 
Toliara; Mahafaly. 

Caridina norvestica Holthuis 1965 
Mahajanga and Lake Mahajamba. 



Caridina petiti Roux 1929 

Known only from type locality near Ambila, 

Madagascar. 



eastern 



Caridina troglophila Holthuis 1965 

Known only from type locality; Ambovonomby Cave, 

Namoroka, north-west Madagascar. 

Caridina xiphias Bouvier 1925 
Several localities. 

Parisia edentata Holthuis 1956 

Antsingy Mountains, near Bekopaka, Mahajanga Province. 

Parisia macrophthalma Holthuis 1956 

Grotte des Fanihy, Ankarana Mountains, north of Ambilobe. 

Parisia microphthalma (Fage 1946) 

Grotte des Fanihy, Ankarana Mountains, north of Ambilobe. 

Typhlopatsa pauliani Holthuis 1956 

Mitoho Cave, north-east corner of Lake Tsimanampetsotsa, 

Mahafaly Province. 

Family PALAEMONIDAE 

Macrobrachium hildebrandti (Hilgendorf, 1893) 
Endemic. 

Macrobrachium petiti (Roux, 1934) 
Endemic. Vatomandry. 

Macrobrachium patsa (Coutiere, 1899) 

(= Macrobrachium lepidactvlus ) 

Patsa river prawn, Bouquet patsa, 'orana', 'camaron'. 

Endemic. Andampy; Manahara River; Onilahy; Bay 

d'Antongil. 

Family POTAMONIDAE (endemic species) 
Gecarcinautes antongilensis antongilensis (Rathbun 1905) 
Baie of Antongil, Toamasina, Ambilobe, Manambato. 

Gecarcinatus antongilensis vondrozi Bott, 1965 
Known only from type locality, Vondrozo (700-800m). 

Gecarcinatus goudoti (Milne-Edwards 1853) 
Antananarivo, Ivoloina River, Chambendiana River. 

Hydrothelphu8a agilis agilis Milne-Edwards 1872 

Several localities and rivers in areas including Sakaleony, 

Antananarivo, Toamasina, Bombetok, Ambodrina stream at 

Perinet, Ambetolamy stream, Ranomafana, Beforona, 

Schambendrama. 

Hydrothelphusa agilis madagascariensis (Milne-Edwards 

1872) 

Antseranana, Mount d'Ambre, Bombetoke, Sakaleony River, 

Antananarivo, Sakavalana, Betampona reserve. 



Appendix 2: faunal lists 

Hydrothelphusa humbloti (Rathbun, 1904) 

Woods. Toamasina, Taolanaro, Andrafialava, Sakalava, 

Ambohitantely (1700 m). 

Madagapotamon humberti Bott. 1965 

Known only from type locality. Woods between Ankara and 

Analamera. 

Madagapotamon gollhardi Bott, 1965 

Known only from type locality. Cave at Ankara. 

Macrobrachium ankaraharae (Nobili, 1906) 
Ankaraha, Antseranana. 

Family PARASTACIDAE 

Astacoides m. madagascarensis (H. Milne Edwards & 
Audouin, 1839) 

Around Antananarivo. This subspecies occurs most 
commonly in the market at Antananarivo. 

Astacoides madagascarensis caldwelli (Bate, 1865) 

Streams on eastern slopes of Ankaratra Mountains, i.e. to 

south and slightly to south-west of m. madagascarensis . 

Astacoides madagascarensis granulimanus Monod & Petit, 

1929 

South-east, distinct from madagascarensis and caldwelli but 

coincides with betsileoensis . 

Astacoides madagascarensis betsileoensis Petit, 1923 

('orambanonga' or 'orambato') 

South: Betsileo, Fianarantsoa, Ikongo Mountains and forest. 

Family SIDIDAE 
Pseudosida bidentata Herrick, 1884 

Latonopsis australis Sars, 1888 

Latonopsis orientalis Sars, incertae sedis 

Diaphanosoma paucispinosum Brehm, 1933 

Family DAPHNIIDAE 
Daphnia carinata King, 1852 

Ceriodaphnia laticaudata P.E. MUller, 1867 

Ceriodaphnia rigaudi Richard. 1894 

Ceriodaphnia c. quadrangula (O.F. Muller, 1785) 

Simocephalus serrulatus (Koch, 1841) 

Family MOINIDAE 
Moina hartwigi Weltner, 1898 

Moina cf. dubia de Guerne & Richard, 1892 likely to be 
conspecific with M. hartwigi 

Moinodaphnia macleavi (King, 1852) 

Family MACROTHRICIDAE 
Echinisca odiosa (Gurney, 1907) 

Echinisca orbicularis (Brehm, 1930) 

Echinisca madagascariensis (Brehm, 1933) 

Echinisca chevreuxi de Guerne & Richard, 1892 



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APPENDIX 3. SPECIES ACCOUNTS 

This appendix contains individual accounts for selected groups of Madagascan animal species, 
most of them threatened or possibly so. Classifications of degree of threat follow the IUCN 
definitions set out below, though it should be noted that in some cases designations are 
preliminary and should not be taken as official IUCN categories. 

The categories are defined as follows: 

Extinct (Ex) 

Species not definitely located in the wild during the past 50 years (criterion as used in The 

Convention on International Trade in Endangered Species of Wild Fauna and Flora - CITES). 

Endangered (E) 

Taxa in danger of extinction and whose survival is unlikely if the causal factors continue 
operating. 

Included are taxa whose numbers have been reduced to a critical level or whose habitats have 
been so drastically reduced that they are deemed to be in immediate danger of extinction. Also 
included are taxa that are possibly already extinct but have definitely been seen in the wild in 
the past 50 years. 

Vulnerable (V) 

Taxa believed likely to move into the Endangered category in the near future if the causal 
factors continue operating. 

Included are taxa of which most or all of the populations are decreasing because of 
over-exploitation, extensive destruction of habitat or other environmental disturbance; taxa 
with populations that have been seriously depleted and whose ultimate security has not yet 
been assured; and taxa with populations which are still abundant but are under threat from 
severe adverse factors throughout their range. 

Rare (R) 

Taxa with small world populations that are not at present Endangered or Vulnerable, but are at 
risk. 

These taxa are usually localized within restricted geographical areas or habitats or are thinly 
scattered over a more extensive range. 

Indeterminate (I) 

Taxa known to be Endangered, Vulnerable, or Rare but where there is not enough information 
to say which of the three categories is appropriate. 

Insufficiently Known (K) 

Taxa that are suspected but not definitely known to belong to any of the above categories, 
because of lack of information. 

Out of Danger (O) 

Taxa formerly included in one of the above categories, but which are now considered relatively 
secure because effective conservation measures have been taken or the previous threat to their 
survival has been removed. 

N.B. In practice, Endangered and Vulnerable categories may include, temporarily, taxa whose 
populations are beginning to recover as a result of remedial action, but whose recovery is 
insufficient to justify their transfer to another category. 



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An environmental profile of Madagascar 



APPENDIX 3. A. BIRDS 



Data sheets for the following species are provided, extracted from: Collar, N.J. and Stuart, S.N. 
(1985) Threatened birds of Africa and related islands: the ICBP/IUCN Bird Red Data Book, 
3rd edition, part 1. ICBP/IUCN, Cambridge. 



K Tachyhaptus pelzelnii 

E Tachybaptus rufolavatus 

K Ardea humbloti 

V Anas bernieri 

E Aythya innotata 

E Haliaeetus vociferoides 

E Eutriorchis astur 

R Mesitornis variegata 

K Mesitornis unicolor 

R Monias benschi 

I Sarothrura watersi 

K Amaurornis olivieri 

R Charadrius thoracicus 

Ex Coua delalandei 



I Tyto soumagnei 

R Brachypteracias leptosomus 

R Brachypteracias squamiger 

R Atelornis crossleyi 

R Uratelornis chimaera 

I Neodrepanis hypoxantha 

R Phyllastrephus apperti 

R Phyllastrephus tenebrosus 

R Phyllastrephus cinereiceps 

R Xenopirostris damii 

I Xenopirostris polleni 

R Monticola bensoni 

I Crossleyia xanthophrys 

I Newtonia fanovanae 



A full reference list is provided at the end of this section. 



MADAGASCAR LITTLE GREBE 



INSUFFICIENTLY KNOWN 



Tachybaptus pelzelnii (Hartlaub, 1861) 



Podicipediformes: Podicipedidae 



SUMMARY This endemic Madagascar waterbird, common and widespread in the recent past, 
is known to have suffered a considerable decline in certain areas and, in view of the variety of 
threats it faces, it is treated here as a case requiring precautionary or preventive measures. 

DISTRIBUTION The Madagascar Little Grebe is endemic to Madagascar where it is 
widespread from sea-level to 1800 m, and absent only from the subdesert region in the south, 
including Lake Tsimanampetsotsa (Delacour 1932a, Rand 1936, Milon et al. 1973), although 
there is a specimen in NHMW from the south-west coast (H. Schifter per Z. J. Karpowicz in 
litt. 1983). 

POPULATION The species was considered common, 1929-1931 (Delacour 1933, Rand 1936), 
and locally common, 1942-1944 (van Someren 1947). In 1973 it was still described as common 
except for at least 15 km around Antananarivo, where it was rare (Milon et al. 1973), but other 
evidence suggests that it was probably no longer common anywhere at that stage and is likely 
to be less so now. Thus it was found to be abundant at Lake Ihotry in the south-west in 
December 1929 (Rand 1936) but was extremely rare there, 1960-1966 (Appert 1971b), though 
100-150 were present on it in August 1983 (O. Langrand in litt. 1984); and, although birds 
were common at Lake Alaotra, 1929-1931 (Delacour 1932a), in a three-month study of grebes 
in north-central Madagascar ranging from south and west of Antananarivo to north of Lake 
Alaotra, 1960, only 10 of this species were seen (at Lake Alaotra and around Andilamena 
30 km to the north) and it was "definitely the rarest" of the three species seen and had 
"considerably decreased in numbers" (Voous and Payne 1965). Despite a report that at least 100 
were present at Lake Itasy and on nearby crater-lakes around 1970, this species along with the 
Madagascar Pond-heron Ardeola idae was then regarded as in complete collapse around 
Antananarivo (Salvan 1972a). The factors apparently causing the decline at Lakes Ihotry and 
Alaotra and around Antananarivo are reportedly widespread in Madagascar (e.g. Salvan 
1970,1972b, Appert 1971b), and it seems likely that the species will have declined everywhere 
and may well now be threatened. That it has generally declined has been confirmed by 
occasional observations spanning the past 15 or so years (D. A. Turner in litt. 1983). 

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Appendix 3: species accounts 

ECOLOGY It inhabits lakes, pools and slow stretches of rivers (Rand 1936), preferably those 
most richly vegetated with aquatic plants and notably the water-lily Nymphaea stellata, 
occurring much less often on vegetation-free water (Appert 1971b). The species is considered 
to be less exclusively piscivorous than either of its congeners in Madagascar, the Little 
Grebe Tachybaptus ruficollis (with which it is often seen: O. Langrand in lilt. 1984) and the 
Alaotra Grebe T. rufolavatus (Voous and Payne 1965, Appert 1971b) (see relevant account); of 
eight stomachs of birds collected, 1929-1931, all held aquatic insects, four also feathers (Rand 
1936); of five stomachs, 1960, feathers were in four, fish in two, insects in two, a crustacean in 
one (Voous and Payne 1965). Breeding appears to occur chiefly at the end of the rainy season 
(February to April), when water-levels are highest and aquatic plants most developed; in 
favourable conditions it evidently also occurs in the austral spring (August to October) (Appert 
1971b; also Rand 1936). In BMNH there is a downy chick from Namoroka (north-west) in 
March and a female ready to lay from Iampasika (south-east) in August (NJC). Clutch-size is 
three to four (Milon et al. 1973); nests may be close to each other (see Appert 1971b). The 
species is forced to move around because many waterbodies annually dry out while others 
shrink greatly in size (Appert 1971b). 

THREATS Apart from the pollution of waters around Antananarivo (Salvan 1970,1972b), two 
major and two less immediately certain threats can be identified, the first three of which are 
interrelated. 

Introduced exotic fish The introduction of herbivorous tilapia into many waterbodies 
throughout Madagascar has apparently resulted in a massive reduction in their vegetation 
(Appert 1971b), e.g. Lake Ihotry had been rich in water-lilies in 1929, but very few were seen 
in 1960-1966 (Appert 1971b) although it was only in October 1960 that the lake was 
successfully stocked with tilapia (Griveaud 1960a). These fish are able to colonise sites away 
from the release area during the rainy season; only very isolated pools or ones which dry out 
every year escape (Appert 1971b). All waters found to hold grebes in north-central 
Madagascar, 1960, had abundant small fish, mainly tilapia (Voous and Payne 1965). The black 
bass Micropterus salmoides is regarded as both a food-competitor and a predator on downy 
young of this and other waterbird species (Salvan 1972a). 

Competition with the Little Grebe The spread through Madagascar of T. ruficollis is outlined in 
Threats under Alaotra Grebe. Its post-1945 increase in abundance appears to be related to the 
conditions created by the introduction of exotic fish, since ruficollis is more piscivorous 
than pelzelnii and occurs widely on vegetation-free waters (Voous and Payne 1965, Appert 
1971b). "As the structural characters of the invading ruficollis more closely resemble pelzelnii 
than rufolavatus, it is not unlikely [see Remarks] that the decline of pelzelnii is caused by the 
recent colonisation of ruficollis. The structure and ecology of these species make it not 
improbable that the decline will continue" (Voous and Payne 1965). 

Hybridisation with the Little Grebe A possible hybrid ruficollis x pelzelnii has been described 
(Benson 1971a) and an apparent pair-bond between birds of these species has been observed in 
the wild (Benson et al. 1976). This evidence, though at present slight, suggests that 
as ruficollis spreads and multiplies while pelzelnii contracts and declines, further interbreeding 
could lead to genetic swamping by the former of the latter. 

Reduction of wetlands Various factors over the past 50 years have resulted, in the Mangoky 
region at least, in less water in rivers and lakes and a lowering of the water-table, so that 
overall there is less grebe habitat (Appert 1971b). Marshes throughout the island have been 
transformed into rice-fields and fish-farms (Salvan 1970,1972b). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED A modern evaluation of the plight of the 
Madagascar Little Grebe is required before appropriate measures can be proposed; nevertheless 
it seems clear that such measures should include the safeguarding of a network of 
vegetation-rich lakes and pools from the introduction of exotic fish. 



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An environmental profile of Madagascar 

REMARKS In the passage (from Voous and Payne 1965) quoted under Threats, the original 
has "likely" for the obviously correct "unlikely": this misprint has been confirmed (K. H. 
Voous in litt. 1983). 



ALAOTRA GREBE ENDANGERED 

Tachybaptus rufolavatus (Delacour, 1932) Podicipediformes: Podicipedidae 

SUMMARY This endemic Madagascar waterbird, known chiefly from Lake Alaotra, is in the 
irreversible process of disappearing through hybridisation with the Little Grebe Tachybaptus 
ruficollis. 

DISTRIBUTION The Alaotra Grebe is known primarily from Lake Alaotra (40 km by 
3-5 km) and adjacent marshes, at c. 700 m in north-eastern Madagascar (Delacour 1932a, 
Lavauden 1937, Voous and Payne 1965). A prediction that it would be reported from Lake 
Itasy and other marshes in central Madagascar (Lavauden 1937) has been partially fulfilled, 
with records from Ankazobe (80 km north of Antananarivo) in December 1947 (Salvan 1971), a 
crater-lake north of Analavory (80 km west of Antananarivo) on an unknown date (but 
apparently around 1970) (Salvan 1972a), "Mianinarivo" (correctly, Miarinarivo: J. T. 
Hardyman in litt. 1984) (one town of this name is near Analavory and just north of Lake Itasy, 
another is 100 km north of Lake Alaotra) on an unknown date (Voous and Payne 1965), and 
Lake Kazanga (just south of Lake Itasy) in July 1971 (when at least 10 were seen) (Salvan 
1972a). Moreover, the species has been collected as far south as the Isalo massif, in January 
1963, and as far west as Mahajanga in November 1969 (see map in Salvan 1971), and it was 
seen between the Antsingy massif and Antsalova (near the coast due west of Antananarivo) in 
July 1970 (Salvan 1971) and in the Antsingy reserve (R.N.I, no. 9 du Tsingy de Bemaraha) 
itself on an unknown date (but apparently around 1970) (Salvan 1972b) (this and the previous 
record may perhaps be the same). However, it is to be observed that, since hybridisation with 
the longer-winged, dispersive Little Grebe Tachybaptus ruficollis has been taking place from at 
least 1929, and had seemingly intensified by 1960 (see under Threats), and since many hybrids 
can be extremely difficult to distinguish as such (see Voous and Payne 1965), the validity of 
many - if not all - of these records away from Lake Alaotra (which remains the only known 
breeding site) must be doubtful. 

POPULATION Numbers are unknown, but certainly very small. At Lake Alaotra in May 
1929, when 15 specimens were first collected, it was found breeding in fair numbers (Delacour 
1932a); in May/July 1960, when 13 more specimens were collected, the estimated total number 
of birds seen at the lake was 50 (Voous and Payne 1965) (this presumably includes the 13 
collected). More recently it has been stated that this species "seems in expansion" (Salvan 
1972a), presumably as much in terms of numbers as of range; however, the records that are 
evidently the basis of this view, apart from the doubt cast on them under Distribution above, 
can be interpreted in much less encouraging ways, e.g. that they only represent the true but 
hitherto unrecognised distribution of the species, or even that they reflect an unprecedented 
dispersal from Lake Alaotra in the face of deteriorating conditions there. However, 12 birds 
were seen on Lake Alaotra in December 1982 (O. Langrand in litt. 1984). 

ECOLOGY Lake Alaotra is a large but shallow water-body, in 1929 fringed with dense 
vegetation (dominated by papyrus and reeds) and dotted with water-lilies (Rand 1936). The 
Alaotra Grebe is almost exclusively piscivorous (Voous and Payne 1965), breeding April to 
June in 1929 (Rand 1936), January to March in 1960 (Voous and Payne 1965). Its short wing is 
considered an indication of highly sedentary behaviour (Voous and Payne 1965), but 
subsequent records away from Lake Alaotra have been seen to call this assumption in question 
(Salvan 1972a). 

THREATS The species is threatened by hybridisation with the far more widespread and 
numerous Little Grebe, and by alteration of habitat in its only known breeding area. Although 
only first noted in any numbers in Madagascar in 1945 (Milon 1946), the Little Grebe was 

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Appendix 3: species accounts 

evidently fairly widespread in the island in the nineteenth century (up to 17 skins in museum 
collections), with the earliest record in 1837 and a breeding record from 1895 (Schlegel and 
Pollen 1868, Hartlaub 1877, Oberholser 1900, Delacour 1933, Milon 1951, Benson 1971a). 
Despite reports that ruficollis disappeared from near Antananarivo around 1955 (Salvan 1972a, 
Milon et al. 1973), it was "by far the commonest species" of grebe at and around Antananarivo 
and Lake Alaotra in 1960 (Voous and Payne 1965) and had also become widespread in the 
Mangoky river region by this time (Appert 1971b). Its post- 1945 increase in abundance 
appears to be related to the conditions created by the introduction of exotic fish, especially 
tilapia, into many lakes and pools throughout Madagascar (Appert 1971b). Hybridisation by 
the Little Grebe with the Alaotra Grebe, though first recognised in the 1960s (Voous and 
Payne 1965), has been recorded at least as far back as the 1920s (the type-specimen and up to 
four others of the original series of 15 appear hybrid) (Voous and Payne 1965), and even a 
specimen from 1862 seems suspect (Benson 1971a). Of 39 grebes collected in north-central 
Madagascar in 1960, 13 were rufolavatus, 13 ruficollis, and 13 hybrids or suspected hybrids of 
the two; although there was a bias towards collecting birds that proved to fall into this last 
category, it seemed likely on this evidence that the pure rufolavatus strain was "doomed to 
vanish" (Voous and Payne 1965). Observations at Lake Alaotra in April 1971 confirmed that 
hybridisation with ruficollis was on a large scale (D. A. Turner in litt. 1983). Tilapia were 
already in Lake Alaotra in 1960 (Voous and Payne 1965), and while this may not have been 
directly injurious to the population of the piscivorous rufolavatus (indeed, if rufolavatus is 
truly "in expansion" this may well be due to tilapia) it may have provided greater attraction to 
the more mobile ruficollis and thus accelerated the rate of genetic swamping, and may equally 
have reduced cover needed by rufolavatus for breeding. By 1972 Lake Alaotra was said to be 
of limited interest only, owing to developments there for rice-growing and fish-farming 
(Salvan 1972b), a view confirmed by recent observations (O. Langrand in litt. 1984). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED Nothing can be done to prevent the 
extermination of the Alaotra Grebe in the wild; however, a survey to assess its present 
condition would be most valuable for, if sufficient numbers of "pure" birds survive, it might be 
feasible to devise a programme of captive propagation for them. 



MADAGASCAR HERON INSUFFICIENTLY KNOWN 

Ardea humbloti Milne Edwards & Grandidier, 1885 Ciconiiformes: Ardeidae 

SUMMARY Mostly at best uncommon, this large but very little known Malagasy waterbird 
was reported in 1973 to have declined alarmingly and to face extinction unless completely 
protected, although it appears still to be safe in parts of the west coast of Madagascar. 

DISTRIBUTION The Madagascar Heron occurs thinly throughout western Madagascar, 
chiefly in coastal and adjacent areas, but apparently rarely in the east. Records of this species 
are relatively few and many appear to involve wandering individuals. Only three breeding 
sites appear to have been found, in the extreme north (locality not specified), in the extreme 
south-west (locality not specified), and on Nosy Manitra off the south-west coast, west of 
Pointe Fenambosy (Pointe Barrow) (Milon et al. 1973), although a specimen in BMNH from 
Lake Ihotry, collected on 8 December 1929, is labelled "breeding" and another in MNHN from 
Toliara, 18 May 1948, had well developed testes (NJC). Other localities from which birds have 
been reported are chiefly in the north-west around Mahajanga, including Mahajanga itself 
(Muddiman 1983), Ampijoroa in the Ankarafantsika area (Milon et al. 1973), Ambato-Boeni 
(Salvan 1970), along the Betsiboka River between Ambato-Boeni and Mahajanga (O. 
Langrand in litt. 1984), and Lake Kinkony (Rand 1936); birds have also been found further 
north on the coast opposite Nosy Be (Rand 1936), and well to the south at Berevo on the 
Tsiribihina River (Bangs 1918), at Lakes Masama and Bemamba near Antsalova (O. 
Langrand in litt. 1984), at Lake Ihotry (Rand 1936) and on a marsh between Lake Ihotry and 
Morombe (Muddiman 1983). In the central part of Madagascar there have been three records 

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An environmental profile of Madagascar 

from Antananarivo (Milon 1949, Milon et al. 1973), two from Lake Itasy (Salvan 1970,1972, 
H. A. W. Payne per K. H. Voous in litl. 1983), one at the lake near Antsimangana, north of 
Lake Alaotra towards Andimalena, 20 June 1960 (H. A. VV. Payne per K. H. Voous in litt. 
1983), and an unspecified number (but more than three) from Lake Alaotra itself (Milon et al. 
1973, H. A. W. Payne per K. H. Voous in litt. 1983). Although the type-specimen was from 
the "east coast" (Milne Edwards and Grandidier 1885), the only other record from the east is of 
an immature that stayed near Maroantsetra from December 1982 to April 1983 (O. Langrand in 
litt. 1984). Individuals have thrice been recorded from the Comoro Islands: Moheli in 
September 1958 (Benson 1960), Mayotte in October 1965 (Forbes-Watson 1969), and again on 
Mayotte in July and August 1974 (D. A. Turner in litt. 1983, A. D. Forbes-Watson pers. comm. 
1984). 

POPULATION Numbers are evidently rather small, and perhaps localised. The colony on 
Nosy Manitra consisted of five to eight nests, July 1948 (Milon 1948); the other colony in the 
south-west consists (or consisted) of "several nests each year" (Milon et al. 1973); at the site in 
the far north it is not clear if more than one nest was found (see Milon et al. 1973). Thirteen 
birds were collected by the Mission Franco-Anglo-Americaine, 1929-1931 (Delacour 1932a), 
which appears to be all or almost all of the birds seen during that period of study (reported as 
three to four opposite Nosy Be, "a few" at Lake Kinkony, four at Lake Ihotry: see Rand 1936). 
All other records appear to concern single individuals only. In 1973 it was reported that recent 
observations had indicated an alarming decline (Milon et al. 1973), but further details were not 
and have not subsequently been given. Despite all this, two independent observers in the 1970s 
and 1980s provide more encouraging information, the species being thought "not uncommon" in 
some areas of the west coast between Mahajanga and Morondava, though rare elsewhere (D. A. 
Turner in litt. 1983, O. Langrand in litt. 1984); it has also been found "very common" in two 
areas, along the Betsiboka River, where 40 were counted between Ambato-Boeni and 
Mahajanga, April 1982, and at Lakes Masama and Bemamba, date unspecified (O. Langrand in 
litt. 1984). 

ECOLOGY The ecology of this species is probably much as other large herons Ardea, although 
the large bill, sombre colouring and observed adroitness in mandibulating prey are considered 
evidence of specialisation for feeding on large mobile fish rather than on a wider range of 
aquatic prey (Hancock and Elliott 1978). Both small and large fish (including a 48 cm eel) are 
recorded as food (Rand 1936, Benson 1960, Forbes-Watson 1969); it feeds in shallow water in 
lakes and along river banks and on the seashore (e.g. on reefs, at fish-weirs and in estuaries), 
and is recorded also from rice-fields (Rand 1936, Benson 1960, Forbes-Watson 1969, O. 
Langrand in litt. 1984). Although apparently solitary, it nests in mixed heronries; at one site 
(in the far north of Madagascar) it has been found nesting at ground level in a 
vegetation-swathed coral hollow (Milon et al. 1973). Breeding (clutch-size three) has been 
reported in July (Milon el al. 1973) and is considered likely (from gonad condition) in 
December (Rand 1936). Natural predators may include the Madagascar Fish Eagle Haliaeetus 
vociferoides, since a bird has been seen to be attacked by one of these raptors, escaping by 
diving under water (Langrand and Meyburg 1984). 

THREATS The species is perhaps naturally uncommon and localised, and very possibly in 
competition with the more numerous Grey Heron Ardea cinerea and Purple Heron A. purpurea. 
The species's large size and relative tameness were considered in 1973 to expose it to risk, 
presumably from native hunters, and it was asserted that, having recently suffered an alarming 
decline, it would soon become extinct unless completely and carefully protected (Milon et al. 
1973). In 1961, however, under Decree no. 61-096, both Grey and Purple Herons - although 
represented by endemic Malagasy subspecies - were classified as harmful animals, a situation 
which still obtained in 1973 (Forbes-Watson and Turner 1973): to the untrained eye the 
Madagascar Heron is so like these species that it cannot have escaped any persecution of them 
that may have been - and perhaps still is - officially encouraged. Ardeid colonies commonly 
suffer exploitation by locals for eggs (O. Langrand in litt. 1984). Rice-growing is reportedly 
beginning to alter Lake Bemamba (O. Langrand in litt. 1984). 

CONSERVATION MEASURES TAKEN None is known; however, in 1948 the wood which 
held the colony on Nosy Manitra was protected through a local taboo (Milon 1948, Milon et al. 
1973). 

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Appendix 3: species accounts 

CONSERVATION MEASURES PROPOSED Studies are needed to determine the number and 
distribution of colonies of this species, its ecological requirements and long-term trends: such 
work might be linked with similar work on the Madagascar Pond-heron Ardeola idae, and on 
the endangered Madagascar Fish Eagle. Complete and active protection for it (including the 
banning of further collection of specimens) and for its breeding sites is essential (Milon et al. 
1973, Hancock and Elliott 1978). The prohibition of the taking by locals of colonial 
waterbirds' and seabirds' eggs would be a great step forward for conservation in Madagascar 
(O. Langrand in litt. 1984). For the need for a general ornithological survey of both west and 
east coast wetlands in Madagascar, and for the possible importance of wetlands around Cap 
St. Andre and of a proposed faunal reserve in the Antsalova region, see Conservation Measures 
Proposed under Madagascar Teal Anas bernieri. 

REMARKS "This and perhaps Swinhoe's Egret [Egretta eulophotes, treated in King 1978-1979] 
may claim to be the two heron species which are most in need of every care and protection if 
they are to survive" (Hancock and Elliott 1978). 



MADAGASCAR TEAL VULNERABLE 

Anas bernieri (Hartlaub, 1860) Anseriformes: Anatidae 

SUMMARY This little known and evidently much persecuted duck, endemic to Madagascar, 
has been recorded from a few sites along the west coast and its total numbers must be very low. 

DISTRIBUTION Apart from an apparently unsubstantiated assertion that it occurs on the east 
coast (Milne Edwards and Grandidier 1885) and a specimen in Grenoble collected by L. 
Lavauden at Lake Alaotra on 5 September 1932 (O. Langrand in litt. 1984), the Madagascar 
Teal is known only from localities close to the western coast of Madagascar, from the far north 
as far south as Lake Ihotry. There are four specimens, dated 1934, from Montagne d'Ambre 
(far north) in MNHN, Paris (SNS). The species was collected in June 1969 at Ambilobe (far 
north-west) (Salvan 1970) and in the last century from the "north-western coast" (Schlegel 
1866), this presumably referring to the undated specimen in RMNH from "Bonbetak Baai", i.e. 
the Baie de Bombetoka at Mahajanga (NJC). A pair was seen in September 1983 on Lake 
Kinkony (O. Langrand in litt. 1984). Two birds were collected in July/August 1930 in the 
western savannas near Maintirano (Delacour 1932a, Rand 1936); one was seen at Bekopaka 
around this time (Delacour 1956), and a possible sight-record of a pair in July 1929 at 
Ankavandra (Rand 1936) would constitute the most inland record for the species (up the 
Manambolo river east of Antsalova), although subsequently the Antsalova region (especially 
Lake Bemamba) was shown to be a major area for it in the 1970s (Salvan 1970,1972b, Scott and 
Lubbock 1974): Lake Bemamba is a shallow saline lake drying up in September/October, when 
the species is thought to disperse either to the Soahanina estuary or to the remaining small 
freshwater pools and lakes in the forests and rice-fields (Scott and Lubbock 1974). The species 
has also been recorded in the last century from around Morondava (Grandidier 1868; two 
specimens in RMNH: NJC) and in 1957 (but apparently not subsequently: see Threats) from 
Lake Ihotry (south-east of Morombe) (Griveaud 1960a). These data confirm (but slightly 
extend) the species's range, anticipated and mapped as from Ambilobe to north of Morombe on 
the basis of apparent habitat requirements within the 500 to 1,500 mm isohyets (see Salvan 
1970 and under Ecology). 

POPULATION Although not considered rare on the west coast in the last century (Milne 
Edwards and Grandidier 1885) it was described as very rare and localised by around 1930 
(Delacour 1932a,b); and although it has more recently been judged probably less rare than 
records suggest (Milon et al. 1973) the only evidence of this is from the Lake Bemamba region, 
where 13 birds were shot in 1970 (Salvan 1970,1972b) and, on Lake Bemamba itself, 81 birds 
were seen (10 pairs on the eastern shore, 61 individuals maximum on the western) and no more 
than 120 estimated for the whole lake, August 1973 (Scott and Lubbock 1974); this 
concentration was considered probably "the largest for hundreds of miles" (Scott and Lubbock 
1974). 

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An environmental profile of Madagascar 

ECOLOGY In the nineteenth century the Madagascar Teal was reported as occurring in small 
flocks on estuaries, marshes or pools (Milne Edwards and Grandidier 1885), but at least in July 
and August the species appears to occur in rather isolated pairs (Salvan 1970, Scott and 
Lubbock 1974; see also records from 1929 and 1930 above). It appears to occur on marshes 
where recent alluvia and pliocene soils mingle, in herbaceous savanna (with Hyparrhenia 
and Heteropogon), mangrove, and dense deciduous forest (Salvan 1970). Birds feed in shallow 
water or on mud at the water's edge, but have not been observed to drink or fly to fresh water 
(Scott and Lubbock 1974). From courtship activities seen in August, birds were expected to 
breed from mid-September; natives reported breeding in November and April, with clutch-size 
variously claimed as 2-4 and 8-10 (Scott and Lubbock 1974). 

THREATS The hunting of waterfowl in Madagascar was, at least until recently, very 
widespread and very intense (Salvan 1970,1972b, Forbes-Watson and Turner 1973). Although 
hunting pressure at Lake Bemamba did not appear to be great in August 1973, there was some 
poaching (Scott and Lubbock 1974) and the area had been recently opened up for hunting by 
the building of an airport at Ambereny (Salvan 1972b), such that by the early 1980s many 
hunters were coming there by private airplane from (e.g.) Mahajanga and Antananarivo (O. 
Langrand in litt. 1984); moreover, locals have reported that they hunt the Madagascar Teal with 
dogs and plunder nests for eggs (Scott and Lubbock 1974). The impact of such depredations 
elsewhere in Madagascar is not known. The importance to the species of habitat free of the 
influence of tilapia and black bass Micropterus salmoides is also unknown, but the absence of 
records from Lake Ihotry after 1957 may indicate that introduced fish pose a threat to the 
species (for details see Threats under Madagascar Little Grebe Tachybaptus pelzelnii). At least 
in the southern part of the Teal's range (in the Mangoky region), various factors over the past 
50 years have resulted in less water in rivers and lakes and a lowering of the water-table, so 
that overall there is less habitat for aquatic birds (see Appert 1971b). Marshes throughout the 
island have been transformed into rice-fields and fish-farms (Salvan 1970,1972b), and 
rice-growing is now reportedly beginning to alter Lake Bemamba (O. Langrand in litt. 1984). 

CONSERVATION MEASURES TAKEN Hunting is supposed to be banned on Lakes Bemamba 
and Masama in the Antsalova region, also on parts of Lakes Kinkony and Ihotry 
(Andriamampianina 1976). The species is listed on Appendix II of CITES, to which 
Madagascar is a party. 

CONSERVATION MEASURES PROPOSED The area west of the north-south line between 
Antsalova and Bekopaka is so rich ornithologically - but particularly because of its population 
of Madagascar Teal - that a faunal reserve there has been urged, if only at least for Lake 
Bemamba (Salvan 1972b; also Salvan 1970), formally recommended (IUCN 1972) and supported 
(Milon et al. 1973, Scott and Lubbock 1974), but no action appears to have been taken; such a 
reserve would form a valuable westwards extension of the existing reserve at Antsingy (R.N.I, 
no. 9 du Tsingy de Bemaraha), and would be likely to provide a major sanctuary for several 
other threatened bird species, notably the Madagascar Fish Eagle Haliaeetus vociferoides, 
Madagascar Heron Ardea humbloti and Sakalava Rail Amaurornis olivieri (see relevant 
accounts), and also perhaps the Madagascar Pond-heron Ardeola idae. Reassessment of the 
Lake Bemamba situation is now urgent, especially given that there were 70% more waterfowl 
in the 1940s than in 1973 (Scott and Lubbock 1974). In general, this species deserves a detailed 
study at one site to determine its annual requirements and a survey throughout western 
Madagascar to determine its remaining populations and strongholds. Some of this work could 
be coupled with attempts to locate populations of the Sakalava Rail, and with survey work on 
the Madagascar Fish Eagle, Heron, and Pond-heron. In this respect it is to be noted that the 
extensive wetlands (as shown in IGNT 1964) that lie between Ankasakasa/Cap St. Andre and 
Tambohorano appear to have been wholly unstudied by ornithologists and merit inclusion in 
any future survey. Moreover, the wetlands and associated shorelines along the east coast, from 
Sambava northwards and Toamasina southwards, have been similarly neglected at least in this 
century, and in view of nineteenth century records from the east for no fewer than four 
threatened "west coast" birds (Madagascar Heron, Teal, Fish Eagle and Plover Charadrius 
thoracicus) and of the likely importance of these wetlands for many other bird species, a 
general ornithological survey is clearly needed along the coastlines indicated above. 



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REMARKS Only one specimen of this duck appears ever to have been kept in captivity; 
proved hardy (Delacour 1956). 



MADAGASCAR POCHARD ENDANGERED 

Aythya innotata (Salvadori, 1894) Anseriformes: Anatidae 

SUMMARY This freshwater diving duck, endemic to Madagascar, is extremely poorly known 
and since 1930 it has become increasingly rare, but nothing appears to have been done to help 
it. 

DISTRIBUTION The Madagascar Pochard is apparently confined to lakes and pools in the 
northern central plateau of Madagascar. The main site for the species is Lake Alaotra 
(Delacour 1932a, b, Rand 1936, Lavauden 1937, Milon et al. 1973), although there have been no 
published records from there since the 1930s. However, two flocks (of five and three bi-ds) 
were seen in the south-east part of the lake between Andreba and Ambatosoratra, 26 May 
1960, a flock of 20 (one shot, now in ZMA) was seen at Ambatosoratra, 9 June 1960, and a 
flock of five was seen on the north-east side near Imerimandroso, 5 July 1960 (H. A. W. 
Payne per K. H. Voous in litt. 1983); but a recent two-week search of Lake Alaotra failed to 
locate the species (O. Langrand in litt. 1984). In the 1930s Lake Itasy (west of Antananarivo) 
was identified as another locality (Lavauden 1937) but there are no subsequent records despite 
visits in 1969-1971, when the single record for the Antananarivo area was of a pair on Lake 
Ambohibao, 18 March 1970 (Salvan 1970,1972a). Around 1930 the species was seen on a small 
pond near Antsirabe (Rand 1936; see Remarks) and it was recently noted that three were 
collected in 1915 at Ambatomainty, near Maevatanana (Benson et al. 1976). On the 15 June 
1960 two were seen at a barrage near Ambadivato, in the Andilamena region 70 km north of 
Lake Alaotra (H. A. W. Payne per K. H. Voous in litt. 1983). The type-specimen is from 
Betsileo country (Warren 1966), i.e. the southernmost named area for the species (Betsileo 
people mapped in Deschamps 1960, also Locamus 1900). 

POPULATION Numbers are probably at best extremely small. Around 1930 the species was 
common and bred at Lake Alaotra, and 27 were collected (Delacour 1932a, b, Rand 1936). The 
lake was revisited several times in the 1930s and live birds were captured (Webb 1936,1954). 
Since then it has become increasingly rare (Milon et al. 1973). Indeed, since this time the only 
published record is of the pair seen in 1970 (see Distribution). Two independent observers in 
Madagascar during the 1970s and 1980s are united in the belief that this bird is on the brink of 
extinction (D. A. Turner in litt. 1983, O. Langrand in litt. 1984). 

ECOLOGY The Madagascar Pochard is (or was) found on lakes, pools and freshwater marshes 
with open water, where it feeds by diving; it is rather solitary, otherwise in pairs, and not easy 
to observe; it nests in a large tuft of reeds or aquatic vegetation, March/April, clutch-size 
being two (Milon et al. 1973). 

THREATS Large-scale duck-shooting has been blamed for the evidently disastrous decline of 
this species (Forbes-Watson and Turner 1973). The introduction of black bass Micropterus 
salmoides and other exotic fish (e.g. tilapia) into the lakes and pools of the high central 
plateaus has certainly had a serious impact on native wildlife (see Salvan 1970) and may be 
responsible for the loss of food and/or destruction of young of this species. Gill-net fishing of 
exotic fish may also take a heavy toll of adults (A. D. Forbes-Watson pers. comm. 1984). By 
1972 Lake Alaotra was said to be of limited interest only, owing to developments there for 
rice-growing and fish-farming (Salvan 1972b), a view confirmed by recent observations (O. 
Langrand in litt. 1984). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED Legal protection for this species (and many 
others endemic to Madagascar) was urged in a letter to the Director, Service des Eaux et 

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Forets, Chasse et Peche, over 10 years ago (see King 1978-1979). It is not known if any 
measures were adopted. A survey is now urgently needed to determine its distribution and 
numbers, and to provide information from which its conservation can be planned and 
implemented. This is one species that ought to be savable through captive breeding (see below). 

REMARKS The species was frequently bred in captivity prior to World War II, but it is not 
known to be currently represented in captive collections (Delacour 1959). The locality 
Antsirabe is assumed (and almost certain) to be that at 19°51'S 47°01'E, not that at either 
17°11'S 45 o 01'E or 13°59'S 49°59'E (in TAW 1980). 



MADAGASCAR FISH EAGLE ENDANGERED 

Haliaeetus vociferoides Desmurs, 1845 Falconiformes: Accipitridae 

SUMMARY This little known Madagascar raptor, now confined to rivers and shorelines of the 

west coast north of Morondava, has declined to a point where it may be close to extinction, yet 

a project first proposed in early 1979 to survey the species and determine its needs has 
consistently failed to receive funding. 

DISTRIBUTION The Madagascar Fish (or Sea) Eagle is confined to the west coast of central 
to northern Madagascar, from Morondava north to Diego Suarez. It was formerly reported 
from the east coast (Grandidier 1867, Hartlaub 1877, Milne Edwards and Grandidier 1879), but 
these records all appear to be repetitions of each other and based on a single somewhat 
insubstantial reference to its occurrence near Toamasina in 1862 (Vinson 1865). However, a 
male was collected on 25 December 1879 at Ampahana (specimen in RMNH: NJC), the only 
locality of this name (in Office of Geography 1955) being at 14°45'S 50°13'E, with an adjacent 
coastal lake of the same name, i.e on the north-east coast between Antalaha and Sambava; 
moreover, five days later the same collector (J. Audebert) obtained another male at 
"Andrimpona" (specimen in RMNH: NJC), this presumably being the "Andempona" that is 
marked as the next village (a few kilometres) north of Ampahana, rather than the "Andempona" 
marked as just north of Sambava (in Locamus 1900). In 1891 it was reported as "all along the 
western coast and on the numerous small islands off the north-west of the mainland" (Sibree 
1891) and this is probably close to the true situation at that time, although evidence of its 
occurrence in the southern half of the west coast is extremely feeble. Four main general 
regions have been identified (although these may merely reflect ornithological activity): Nosy 
Be and the coastline opposite, the Lake Kinkony region, the Antsalova region, and the coastline 
between the Mangoky and Fiherenana Rivers, the species apparently being extinct now in this 
last region. In the first of these regions, eight specimens were collected in two weeks around 
1930 on the mainland opposite Nosy Be (Rand 1936) and there are recent reports of the species 
from Nosy Be itself (Thiollay and Meyburg 1981, D. A. Turner in litt. 1983). In the second 
region, there are records from Mahajanga (Kaudern 1922), Lake Kinkony itself, Ambararatabe 
and Soalala (Rand 1936); in August 1969 the area in the Soalala - Namakia - Lake Kinkony 
triangle was identified as a major stronghold, at least 1 1 birds being seen in three days between 
Mahajanga and Lake Kinkony (D. A. Turner in litt. 1983), and a pair was seen there in 1980, 
east of Mitsinjo along the Mahavavy River (Thiollay and Meyburg 1981). In the third region, 
eight birds were seen over Lake Masama and the Manambolo River in July 1970 (Salvan 1971, 
Milon et al. 1973, Langrand and Meyburg 1984) and there have been more recent records 
(Thiollay and Meyburg 1981), including four adults and two juveniles over Lake Masama in 
June 1982 (Langrand and Meyburg 1984), so that the rectangle of the lakes and marshes 
between Antsalova, Bekopaka and the sea is now regarded as the last likely area offering hope 
for the species's survival (Meyburg 1979a, Langrand and Meyburg 1984). In the fourth region, 
the species was reported from near Morombe around 1930 (Rand 1936) and as frequent in one 
area around 1960, but not to be found a decade later (Milon et al. 1973); there were in fact 
seven sightings of single birds in the Morombe region, 1959-1975 (Langrand and Meyburg 
1984). It is probable that the species was recorded at several unnamed sites along the 
north-west coast around 1930, given that 27 specimens were collected there "from west of 
Montagne d'Ambre" (specimen in BMNH: NJC) "to Lake Kinkony" (Delacour 1932a); it was 

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Appendix 3: species accounts 

reported near Antsohihy in the 1940s (van Someren 1947), and there are specimens in BMNH 
and MNHN from Anorontsangana, north of Maromandia (NJC,SNS). Breeding was reported in 
the early 1970s from Lake Ampijoroa (Ankarafantsika), well inland from Mahajanga (Salvan 
1971, Milon et al. 1973), but the pair involved was reported not to have produced young for 
several years prior to 1978 (Meyburg 1979a, B.-U. Meyburg pers. comm. 1983; see Remarks 
under Van Dam's Vanga Xenopirostris damii). Nesting has also recently been recorded on a 
small island c. 30 km west of Diego Suarez, and there is a recent record from north of 
Maintirano, five birds being reported shot in this region (Langrand and Meyburg 1984). There 
appear to be two or three old records from Mauritius (Benson 1970). 

POPULATION In the last century the species was not rare and was often seen in the 
north-west (Schlegel and Pollen 1868), was still fairly common there around 1930 (Delacour 
1932a, Rand 1936) but was considered scarce in the 1940s (van Someren 1947). Despite the 
fairly recent records from Lakes Kinkony and Masama (see above), at the end of the 1970s it 
was estimated that only 10 pairs survived (Meyburg 1979a, Thiollay and Meyburg 1981). More 
recently, this estimate has been raised to 30 pairs (O. Langrand in litt. 1984). Nevertheless, the 
species is still to be considered one of the rarest birds of prey in the world (Langrand and 
Meyburg 1984). 

ECOLOGY It is largely a coastal species, inhabiting estuaries and mangrove-bordered bays 
where shallow waters facilitate fishing, but also lakes and rivers (Grandidier 1867, Schlegel and 
Pollen 1868, Rand 1936). It takes fish from water in a plunge-dive (Grandidier 1867, Milne 
Edwards and Grandidier 1879, Rand 1936), though attacks on large waterbirds 
(Spoonbill Platalea alba and Madagascar Heron Ardea humbloti) have been witnessed 
(Langrand and Meyburg 1984). It is commonly found in pairs at traditional sites (Grandidier 
1867, Rand 1936), and builds a large nest in the highest tree of forest along the coast or up a 
river (Schlegel and Pollen 1868), though the nest near Diego Suarez (see Distribution) was on a 
cliff 6-8 m high (Langrand and Meyburg 1984). It breeds in the dry season (Milon et al. 
1973), towards the start of the rains (Milne Edwards and Grandidier 1879), but not in 
November/December (Rand 1936). Only one young is raised (Milne Edwards and Grandidier 
1879, Milon et al. 1973) though two eggs are laid (Milon et al. 1973, Langrand and Meyburg 
1984). Age of first breeding is put at four or five years (Milon et al. 1973). The records from 
Mauritius (and also perhaps from the east coast) suggest a powerful dispersive ability. 

THREATS The reasons for the decline of this species are unclear (Langrand and Meyburg 
1984). Shooting by amateur hunters was suspected to have caused its disappearance between 
the Mangoky and Fiherenana Rivers (Milon et al. 1973), and five birds have been reported shot 
in recent years in the Maintirano area (Langrand and Meyburg 1984); deliberate destruction of 
nests is also stated to occur (Thiollay and Meyburg 1981). Rice-growing is reportedly 
beginning to alter Lake Bemamba (O. Langrand in litt. 1984). 

CONSERVATION MEASURES TAKEN A leaflet has been produced to increase public 
awareness of the species's plight (Fonds d' Intervention pour les Rapaces no. 9 [1983]: 44, 
Langrand and Meyburg 1984). Along with all Falconiformes, it is included on Appendix II of 
CITES, to which Madagascar is a party. 

CONSERVATION MEASURES PROPOSED Full protection for this bird is merited (Milon et 
al. 1973). A faunal reserve has been urged for the Antsalova region, identified above (under 
Distribution) as perhaps this species's last stronghold (see Conservation Measures Proposed 
under Madagascar Teal Anas bernieri). Since early 1979, a proposal to survey and census it 
from the air, as a first step to determining further conservation action, has languished for lack 
of financial support, despite repeated inclusion in the annual ICBP programme. It is to be 
noted that a similar problem exists for the Madagascar Serpent Eagle Eutriorchis astur and that 
these two raptors, among the world's rarest and yet without any conservation action on their 
behalf, remain the highest priorities for such action at present (Langrand and Meyburg 1984). 
For the need for a general ornithological survey of east coast wetlands in Madagascar, and for 
the possible importance of wetlands around Cap St.Andre, see Conservation Measures Proposed 
under Madagascar Teal. 



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REMARKS It is to be hoped that in the course of the proposed survey and resulting research 
and action for this species it will be possible to accommodate the study and conservation of 
three other birds of considerable importance, the Madagascar Teal, the Madagascar 
Heron Ardea humbloti (see relevant accounts) and the Madagascar Pond-heron Ardeola idae. 



MADAGASCAR SERPENT EAGLE ENDANGERED 

Eutriorchis astur Sharpe, 1875 Falconiformes: Accipitridae 

SUMMARY This very poorly known Madagascar rainforest raptor was last seen by an 
ornithologist over 50 years ago, and hopes for its survival are largely pinned on the 
conservation of adequate areas of primary forest in the central-east and north-east of the island. 

DISTRIBUTION The Madagascar Serpent Eagle is confined to the eastern rainforests of 
Madagascar, and known from only eight specimens, all collected more than 50 years ago (four 
in MNHN, two in AMNH, one in BMNH and one in Grenoble) (A. Fayaud in litt. 1983, G. S. 
Keith in litt. 1983, NJC); a further specimen reportedly in Berlin (Lavauden 1937) cannot be 
traced (B.-U. Meyburg in litt. 1984). The species was first described from a single specimen 
collected (presumably around 1874) "in the southern portion of Madagascar" (Sharpe 1875) 
though the locality was later identified as "Ampasimanavy", a hamlet in the forest a day's 
march from Andakana village, in the Mangoro valley between Antananarivo and Mahanoro 
(Milne Edwards and Grandidier 1879; see Remarks). A second bird, dated 1883 and labelled 
simply "Madagascar", was collected by L. Humblot (specimen in MNHN: NJC). In April 1924 a 
male was obtained in forest at Fito, i.e. Sihanaka forest (specimen in AMNH: G. S. Keith in 
litt. 1983). The Expedition Citroenen en Afrique obtained a bird at an unknown date and from 
an unknown locality (specimen in MNHN: NJC), although it is known that the Citroen team 
arrived in Antananarivo in June 1925 (R. D. Etchecopar in litt. 1984). Four specimens were 
collected in the period 1928-1930, one from Rogez at 900 m in eastern central Madagascar 
(18°50'S 48°35'E), December 1928 (Lavauden 1932, Benson et al. 1976), one from 
Analamazaotra near Perinet (i.e. also near Rogez), 11 June 1930 (specimen in Grenoble: A. 
Fayaud in litt. 1983), and two from around Maroantsetra (one at sea level at Bevato, 40 km 
north-west of Maroantsetra up the Vohemar River, 8 May 1930, the other at 600 m at 
"Ambohimarahavary" [see Remarks under Short-legged Ground-roller Br achy pier acias 
leptosomus], two days' march north-east of Maroantsetra, 6 July 1930) in the north-east of the 
island (Rand 1932,1936). The species has been reported to occur as far south as Farafangana 
(Lavauden 1937), although there appears to be no evidence for this other than that a bird, 
either this species or Henst's Goshawk Accipiter henstii (see Remarks), was seen at Vondrozo 
(inland from Farafangana), June or July 1929 (Rand 1932). A forestry official reported 
making four or five sightings of a raptor closely answering this species's description over the 
period 1964-1977 in the Marojejy Reserve, north-west of Andapa in north-eastern Madagascar 
(Meyburg and Meyburg 1978, Meyburg 1979b, Thiollay and Meyburg 1981). There have been 
no other reports since 1930 though it is hoped the species may also survive on the Masoala 
peninsula in the north-east (Meyburg and Meyburg 1978, Meyburg 1979b). 

POPULATION Numbers are unknown, but the species was repeatedly described as very rare 
fifty years ago (Delacour 1932a, Lavauden 1932, Rand 1936), so presumably it is very much 
more so at present: indeed it is authoritatively considered one of the six rarest birds of prey in 
the world (Langrand and Meyburg 1984). However, since it has also been said to be very shy 
(Lavauden 1932) it has conceivably avoided detection in several areas, although the forestry 
official who claimed to have seen it in the Marojejy Reserve (see above) considered it 
relatively fearless (B.-U. Meyburg in litt. 1983). At any rate, to treat the species as extinct 
(Day 1981) is on present information irresponsibly pessimistic. 

ECOLOGY This bird inhabits primary rainforest, although it has also been recorded in 
secondary growth at the edge of dense forest (Lavauden 1932, Rand 1936). Its short wings and 
long tail are considered adaptations for flight below the canopy (Lavauden 1932,1937), 
although it is also considered a bird of the tree-tops (Lavauden 1937). One of the birds 

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Appendix 3: species accounts 

collected near Maroantsetra contained part of a very large chameleon (Rand 1936), but the 
species is also reported to attack lemurs and even poultry belonging to forest guards (Lavauden 
1932) and to feed chiefly on mammals (Lavauden 1937). There appears to be no direct 
evidence that it eats snakes (see Remarks). There are no breeding data (Lavauden 1937). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar. "The present wholesale 
destruction of the forest" (i.e. rainforest) was being lamented almost 100 years ago (Baron 1890) 
but has continued unabated throughout the present century (Humbert 1927, Heim 1935, Rand 
1936, Swingle 1937, Louvel 1950, Chauvet 1972, McNulty 1975, Guillaumet 1981) and is now 
proceeding so "incredibly fast" that "good places four or five years ago are already destroyed" 
and "within the next five years ... all the good [i.e. rich, lowland] forests will vanish" (B.-U. 
Meyburg in litt. 1983). It is estimated that in the years 1981-1985 loss of primary forest in 
Madagascar will be 35,000 ha per year, most of this in the eastern rainforests and most of it as 
a result of slash-and-burn ("tavy") cultivation. The de-gazetting of the Masoala Forest Nature 
Reserve (R.N.I, no. 2) is highly regrettable (see Conservation Measures Proposed). 

CONSERVATION MEASURES TAKEN The species's reported presence in the Marcjejy 
Reserve (R.N.I, no. 12), which covers 60,150 ha (Andriamampianina 1981), reinforces the 
importance of this protected area; however, it has been pointed out that only the lower parts of 
the reserve provide suitable habitat, the higher-lying areas lacking sufficient vegetation 
(Meyburg 1979b). A "Special Reserve" also exists at Perinet-Analamazaotra, where the 
Madagascar Serpent Eagle was once recorded (see Distribution), but only covers 810 ha 
(Andriamampianina 1981) and the species evidently does not now occur there. Along with all 
Falconiformes it is included on Appendix II of CITES, to which Madagascar is a party. 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar: this was formally recommended in 1970 
(IUCN 1972). Complete protection of the intact parts of "Sihanaka forest" is of extreme 
importance, being the single most important tract of unprotected bird habitat at present known 
in Madagascar: with the reasonable exception of the Snail-eating Coua Coua delalandei and the 
Red-tailed Newtonia Newtonia fanovanae (see relevant accounts), all Madagascar rainforest 
birds here treated as threatened have been recorded there, namely the Brown Mesite Mesitornis 
unicolor, Madagascar Red Owl Tyto soumagnei, Short-legged Ground-roller, Scaly 
Ground-roller Brachypteracias squamiger, Rufous-headed Ground-roller Atelornis crossleyi, 
Yellow-bellied Sunbird-asity Neodrepanis hypoxantha, Dusky Greenbul Phyllastrephus 
tenebrosus, Grey-crowned Greenbul P. cinereiceps, Pollen's Vanga Xenopirostris polleni and 
Madagascar Yellowbrow Crossleyia xanthophrys (see relevant accounts). "Sihanaka forest" is 
technically a misnomer, since the Sihanaka people are to the west of the central rainforest belt, 
which is inhabited by the Betsimisaraka people (J. T. Hardyman in litt. 1984); the name appears 
to have been imposed by explorers to stand crudely for the broad belt of humid forest from the 
coast to the Mangoro valley, east and south of Lake Alaotra and in particular in the Toamasina 
hinterland, notably between the towns of Didy and Fito (see, e.g., the map in Delacour 1932a). 
Proposals for a comprehensive ornithological survey of Madagascar's rainforests, to feature 
studies of the Sihanaka forest, the adjacent Zahamena Nature Reserve (R.N.I, no. 3), and other 
protected areas of rainforest, with particular emphasis on the Serpent Eagle, are to be drawn 
up as part of an overall plan for bird conservation and research on the island. A proposal in 
1979 to search for this species in the Marojejy Nature Reserve and later on the Masoala 
peninsula (Meyburg 1979b) was adopted as WWF Project 1368, and the required sums were 
raised; however these sums were not released and the project did not proceed (Langrand and 
Meyburg 1984). It is to be noted that a similar problem has existed for the Madagascar Fish 
Eagle Haliaeetus vociferoides and that these two raptors, among the world's six rarest and yet 
without any conservation action on their behalf, remain the highest priorities for such action at 
present (Langrand and Meyburg 1984). The re-gazetting of the Masoala Forest Nature Reserve 
(R.N.I, no. 2) was formally recommended in 1970 (IUCN 1972). 

REMARKS This species is the only one in its genus (see Sharpe 1875). Concerning the 
type-locality, Andakana is at 19°22'S 48°05'E on the Mangoro River (Office of Geography 
1955, IGNT 1964); neither "Ampasimanavy" nor "Ampasmonhavo" (the name given apropos 

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An environmental profile of Madagascar 

other species in Sharpe 1875) can be traced (Office of Geography 1955, IGNT 1964), but there 
is an "Ampasimaneva" a few kilometres to the south of Andakana (see IGNT 1964) which must 
surely be the site (19°24'S 48°04'E). This is also the type-locality of the Rufous-headed 
Ground-roller (Sharpe 1875) and it is therefore of considerable importance to establish whether 
good forest still stands in that part of the Mangoro valley. Concerning the name "serpent 
eagle", confusion may arise in field studies since one French name for the Madagascar 
Harrier-hawk Polyboroides radiatus is "serpentaire" (A. D. Forbes-Watson pers. comm. 1984): 
possibly "crested eagle" or "forest eagle" would be a more appropriate name for E. astur. It has 
been remarked that there is great similarity between specimens of the Madagascar Serpent 
Eagle and those of Henst's Goshawk (A. Fayard in litt. 1983), and the AMNH specimen from 
Fito was originally labelled as the latter species (G. S. Keith in litt. 1983): given the importance 
of museum material in clarifying the range of the former, a check needs to be made of skins of 
Henst's Goshawk to confirm their identity, and details of any Serpent Eagles thus (or 
otherwise) discovered are requested to be forwarded to ICBP. Not having been seen with 
certainty in the wild for over 50 years, by CITES criteria this species would now be considered 
extinct. 



WHITE-BREASTED MESITE RARE 

Mesitornis variegata (I. Geoff roy Sainte-Hilaire, 1838) Gruiformes: Mesitornithidae 

SUMMARY This rail-like terrestrial forest bird is currently known from only two sites in 
Madagascar, one of which is, however, a protected area. 

DISTRIBUTION Although the White-breasted Mesite was first found in 1834 at an 
unspecified locality in Madagascar, almost a century passed (during which all records of this 
species are attributable to the Brown Mesite Mesitornis unicolor. see Lavauden 1931) before it 
was rediscovered: an adult female was collected on 12 July 1929 in Ankarafantsika forest 
(110 km south-east of Mahajanga), north-west Madagascar, and a nest with two eggs was 
found there in October that year (Lavauden 1931,1932). A year later, on 10-11 November 
1930, two males and a gravid female were collected at Ankarana cliffs, 25 km south-west of 
Tsarakibany, in the far north of the island (Rand 1936). A few were seen in 1971 at 
Ankarafantsika (Forbes- Watson et al. 1973) and further visits there through the 1970s 
consistently resulted in sightings (D. A. Turner pers. comm. 1983), but there appear to be no 
other records for this species. Nevertheless it has been speculated that birds may occur in the 
region between the two known localities, "notably in the Analalava and in the Haut-Sombirano 
[sic]" (Lavauden 1932), and that the Betsiboka River may mark the southern boundary of its 
distribution (Lavauden 1937). The statement that it occurs "in all western Madagascar" 
(Milon et al. 1973) is patently unsubstantiated. However, recently published information 
records the species from north-east of Morondava (specifically: 10 km south-west of 
Marofandilia, 15 km north-north-west and 9 km south of Beroboka and 3 km south of 
Ampamanmrika lake), several hundred kilometres south of the only other site (Appert 1985). 

POPULATION Observations through the 1970s suggest that the species is common at 
Ankarafantsika (D. A. Turner pers. comm. 1983). 

ECOLOGY At Ankarafantsika the species is a ground-dweller in dry forest (Lavauden 1932), 
likewise at Ankarana cliffs, where a pair was found "running about together in rather low dry 
forest, somewhat clear of underbrush" (Rand 1936). Food probably consists of insects and fruit 
(Rand 1951); birds live in pairs on the ground, walking or running with frequent stops and 
changes of direction, but flying poorly (only if threatened by a predator) (Lavauden 1931, 
Rand 1936); the nest is placed low in a bush (60-80 cm above ground), evidently 
October/November (Lavauden 1932, Rand 1936). An association appears to exist between this 
species and the Rufous Vanga Schetba rufa, exactly as for the Subdesert Mesite Monias benschi 



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Appendix 3: species accounts 

(see relevant account) and Lafresnaye's Vanga Xenopirostris xenopirostris (A. D. 
Forbes-Watson pers. comm. 1984). 

THREATS The highly restricted range of this species must be a source of permanent concern 
and vigilance for its welfare. Deforestation is likely to have affected many areas where it 
might have been searched for in north-west Madagascar. Introduced rats, widespread in the 
eastern forests in the 1930s and presumably therefore present in the west, may affect the bird 
adversely (see under Brown Mesite). 

CONSERVATION MEASURES TAKEN The only area where it is currently known to occur 
falls within the Ankarafantsika Nature Reserve (R.N.I, no. 7) (see Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED A study of the status and ecology of this bird at 
Ankarafantsika would help determine where else it might be searched for and what 
management it might require. Ankarana cliffs merit revisiting and careful survey. All such 
work should be undertaken in conjunction with studies recommended under Conservation 
Measures Proposed for Van Dam's Vanga Xenopirostris damii. 

REMARKS The importance of the Ankarafantsika Nature Reserve as the only locality 
currently known for this species and Van Dam's Vanga cannot be overstated. 



BROWN MESITE INSUFFICIENTLY KNOWN 

Mesitornis unicolor (Desmurs, 1845) Gruiformes: Mesitornithidae 

SUMMARY This cryptic and retiring terrestrial rail-like bird of Madagascar rainforest 
apparently possesses a much wider distribution than has previously been appreciated, but may 
be at risk from both forest destruction and introduced mammalian predators. 

DISTRIBUTION The Brown Mesite evidently occurs throughout much of eastern Madagascar, 
although most records are from the circle whose diameter lies between Antananarivo and 
Toamasina. One usually reliable authority gave its range as from Vohimarina (high north-east) 
to Farafangana (south-east) (Lavauden 1932) but there appear to be no records to support the 
choice of these extremes and indeed the same authority later speculated whether the species 
reached even as far south as Mananjary (Lavauden 1937). Reports of the bird from the 
"north-east" (Humblot 1882), the Masoala peninsula (Lavauden 1937) and south of Maroantsetra 
(Lavauden 1932,1937), though in themselves too vague to be regarded with confidence, are 
supported by specimens collected by J. Audebert at Mananara (Antongil Bay), 17 August 1876, 
"Savary" in February and April 1878 and "Maintinbato" in May 1878 (specimens in RMNH: 
NJC; also Fisher 1981): "Savary" cannot be traced (e.g. in Office of Geography 1955, IGNT 
1964) but a letter from the collector to H. Schlegel, dated 4 March 1878, is headed "Savary, 
Antongil Bay, west of Mananara, Ancay border, seven days' journey into the interior" (G. F. 
Mees in litt. 1983), which clearly suggests that the "Maintinbato" (i.e. Maintimbato) in question 
is that just south of Rantabe on the shore of Antongil Bay. The type-specimen was described 
as from the "north-east" (Delacour 1932a) but this was later refined to "around Tamatave" 
(Lavauden 1937). There is a specimen in MRAC labelled as from "Brickaville district", 
February 1928 (NJC). The species occurs in the Sihanaka forest, where four birds were taken 
in 1925, three in April, one in November (specimens in SMF: NJC), where an adult female was 
collected in May 1930 (Lavauden 1932) and whence six further specimens were obtained by 
purchase around this time (Delacour 1932a, Rand 1936). The species is known from the forest 
between Rogez and Fito (Lavauden 1937), was seen at Perinet in 1939 or 1940 (Webb 1954), 
and collected in "Lakato forest" in 1924 (two specimens in MRAC: NJC). Four further 
specimens (in MRAC, SMF and RMNH) are from "Vohibazaha forest, Anivorana district", 
October 1923 (two) and "Marovato", November 1922 and March 1923 (NJC): Vohibazaha, at 
18°48'S 48°33'E, is close to Perinet and Rogez, while of at least 34 localities named "Marovato" 
in Madagascar (see Office of Geography 1955) three, at 18°57'S 48°49'E, 18°41'S 48°36'E, and 
18°27'S 48°41'E, all lie within the general area of forest between Antananarivo and 

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Toamasina. The species was collected on the "south-east coast" around 1876 (Bartlett 
1877,1879), and this otherwise anomalous record was vindicated when nesting birds were found 
at "Bemangidy" north of Taolanaro (Rand 1951; see Remarks). It is to be observed that the 
taboo on this species (see Conservation Measures Taken) extended even to speaking its name 
(Lavauden 1931), so that its existence may often have remained unreported to explorers in 
certain areas; elsewhere, where no taboo applied, its existence had gone undetected even by 
natives (Rand 1951). For these reasons, the assertion that the species did not occur at 
Fanovana (Rand 1936), which may have compounded the judgement that it is highly localised 
in distribution (e.g. Rand 1951, King 1978-1979), is open to doubt (although the forest at 
Fanovana is now all cleared - see under Red-tailed Newtonia Newtonia fanovanae); and on 
present evidence it would seem very possible that the bird may be found at many other 
localities to the north of Toamasina or to the south of Lakato. 

POPULATION The species was not considered rare in the last century (Milne Edwards and 
Grandidier 1885) and in Sihanaka forest it is apparently not very rare (Lavauden 1932). Its 
wariness and keen senses have been likened to those of pittas (Pittidae) so that it "may be 
common without being seen" (Webb 1954); nevertheless, it is recently reported as very scarce 
throughout its range (D. A. Turner pers. comm. 1983). 

ECOLOGY The Brown Mesite inhabits the floor of the thickest and remotest parts of 
rainforest, slipping swiftly on foot through thick vegetation (Lavauden 1931,1932). A bird 
observed by a seated observer "alternately ran rapidly and then remained motionless, its colours 
so harmonizing with the background that it was exceedingly difficult to see when stationary" 
(Webb 1954). Food is probably insects and fruit (Rand 1951); in another account "insects, ants" 
are mentioned (Milne Edwards and Grandidier 1885). The species flies poorly (only if 
threatened by a predator) (Lavauden 1931). Both nests found in the south-east in 1948 (on 24 
November and 25 December) were in rainforest where a thin cover of shrubs and a few herbs 
grew below the trees; both were low (1 and 2 m above ground) in the fork of a sloping tree 
which had lower branches possibly used by the bird to hop up from below; both held one egg, 
and in both cases the incubating female was caught by hand (Rand 1951). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). The hilly country in the south-east where nesting was proven 
in 1948 was evidently in the process of being cleared of forest (see Rand 1951). The brown 
rat Rattus norvegicus and black rat R. rattus may affect mesites adversely (Forbes-Watson and 
Turner 1973), and attention has been drawn to the observation, dating from around 1940, that 
"the eastern forests are now swarming with them, even in the most isolated regions where the 
precipitous nature of the country is unfavourable to human habitation" (Webb 1954). It is also 
speculated whether competition from the Madagascar Wood-rail Canirallus kioloides affects 
the species (D. A. Turner pers. comm. 1983). 

CONSERVATION MEASURES TAKEN The species has been recorded from the area now 
established as the Perinet-Analamazaotra Special Reserve, which covers 810 ha 
(Andriamampianina 1981). The strong taboo amongst the Malagasy people in the central part 
of the eastern forests was based on the fact or belief that when the young are captured the 
adult follows the hunter right back into the village, exhibiting parental concern so like that of a 
human being as to render the species sacred (Milne Edwards and Grandidier 1885); it is 
considered that such a taboo must have helped conserve the bird, at least in the past 
(Forbes-Watson and Turner 1973), and indeed at Perinet the taboo still persists (O. 
Langrand per A. D. Forbes-Watson pers. comm. 1984). 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. 



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REMARKS The locality of the two nests found to date was given as "Bemangidy, Poste 
Mananteina, Fort Dauphin (=TaoIanaro) district" with the addition that "Bemangidy is 72 
kilometres north of Fort Dauphin and is about five miles west of the Indian Ocean" (Rand 
1951). However, the correct names appear to be "Bemangily" and "Manantenina" and the 
correct distances 55 km and 5 km respectively (see IGNT 1964). It should also be noted that 
the view, first aired in the original description (Desmurs 1845), that the Brown Mesite might 
be or was only the female of the White-breasted Mesite Mesi'.ornis variegata has resulted in 
considerable confusion; virtually everything written about the latter in Milne Edwards and 
Grandidier (1885) does in fact refer to the Brown Mesite; the view that two species were 
involved was accepted by Hartlaub (1877) and entertained by Lowe (1924) before being 
confirmed by Lavauden (1931,1932,1937). 



SUBDESERT MESITE RARE 

Monias benschi Oustalet and G. Grandidier, 1903 Gruiformes: Mesitornithidae 

SUMMARY This rail-like terrestrial bird of restricted range within the subdesert region of 
south-west Madagascar, although numerically safe at present, appears to enjoy no protection 
whatever. It is of exceptional biological interest. 

DISTRIBUTION The Subdesert Mesite is restricted to a coastal strip roughly 70 km wide 
between the Mangoky and Fierenana Rivers, south-west Madagascar, ranging from sea-level to 
130 m (Lavauden 1937, Appert 1968, Milon et al. 1973). Within this area its distribution was 
thought "extremely local" (Rand 1936) but other evidence suggests it is widespread (Appert 
1968, Turner 1981). Nevertheless it has not been found north of the Mangoky, despite 
apparently suitable habitat (Appert 1968), and there is no evidence of its occurrence south of 
the Fierenana, despite records at and near Toliara (Hartert 1912, Bangs 1918): the 
type-specimen is from Vorondreo, "25 km east of Tulear (=Toliara)" (Oustalet and Grandidier 
1903), but this locality proves to be on the north bank of the Fierenana (i.e. north-east of 
Toliara) at 23°17'S 43°51'E (Office of Geography 1955). The limit of its range inland up the 
Fierenana has been given as Fativolo (Lavauden and Poisson 1929), at 23°02'S 44°10'E (in 
Office of Geography 1955). 

POPULATION The species has been reported as common and at times abundant over much of 
its range (Turner 1981), but the experience of a very recent observer was much less 
encouraging, though birds were "rather common" at Ihotry village in September 1983 (O. 
Langrand in litt. 1984). 

ECOLOGY The Subdesert Mesite is a ground-dwelling bird, reasonably catholic in choice of 
habitat, primarily requiring areas with dense leaf-litter, at least in patches: thus it is found in 
both sparse and dense brush woodland with or without Didierea, and in open sandy scrub with 
isolated trees and bushes, etc., but it avoids shadeless areas and those where vegetation is so 
close to the ground that passage is obstructed (Rand 1936, Appert 1968). It feeds with 
occasional pecks as it walks along, but mainly by digging in leaf-covered soil (Appert 1968). 
Stomachs have been found to contain caterpillars, beetles, millipedes, cockroaches, 
grasshoppers, seeds, and pieces of shell and sand (Lavauden and Poisson 1929, Rand 1936, 
Appert 1968; also specimen-labels in MNHN: NJC). Parts of certain orchids are reported by 
natives to be favoured, and damage to orchids has been noted (Appert 1968). Birds are 
gregarious, generally in groups of four to six, occasionally up to ten, rarely alone; two together 
always represents a pair, at whatever season (Appert 1968). A report of groups up to 30-40 
(Lavauden 1931) has apparently not been corroborated. "Territorial fighting" has been 
witnessed (Steinbacher 1977), but it is unclear if birds are group-territorial. Females are 
bolder than males (Rand 1936, Appert 1968). Although in one set of observations males were 
found to predominate numerically, and this was cited in support of the species possibly being 
polyandrous (Rand 1936), lengthier field study established no rule in the sexual composition of 
groups (Appert 1968). On the basis of a male and two females with a nest with two eggs, an 
instance of polygyny was assumed (Appert 1968), but this conclusion - though perhaps correct 

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- does not take consideration of other possibilities. Nests (one or two eggs) are placed 1-2 m 
up in trees or on broken-off tree-trunks, accessible without need of flight (Lavauden 1931, 
Rand 1936, Appert 1968). Males were reported by natives to incubate and care for the young 
(Rand 1936) and observations have partially supported this (Rand 1936, Appert 1968), but a 
female has been found incubating and a pair seen feeding young, though with the female 
playing more the role of lookout (Appert 1968). Nesting seems mainly to occur within the 
period of spring rains, October to December, but it may occur earlier or later and two young 
were even obtained in June, in the middle of the extended dry period (Lavauden 1932, Rand 
1936, Appert 1968). The species has been stated not to fly (Delacour 1932a) but it was 
reported to do so at the sound of a dog barking (Lavauden 1931) and there are two recent and 
very similar eye-witness accounts (Appert 1968, Turner 1981); moreover, in structure this bird 
is more adapted for flight and life in trees than the other two mesites (Lowe 1924). An 
association appears to exist between this species and Lafresnaye's Vanga Xenopirostris 
xenopirostris, since birds of the latter species are often found above parties of the former: the 
Mesites possibly flush insect prey for the Vangas and benefit in turn from the Vangas' greater 
vigilance (A. D. Forbes- Watson pers. comm. 1984); for a similar association between a vanga 
and a mesite, see Ecology under White-breasted Mesite Mesitornis variegata. 

THREATS The restricted range of this species must be a source of permanent concern and 
vigilance for its welfare. The Subdesert Mesite shares an identical range with the Long-tailed 
Ground- roller Uratelomis chimaera and occupies the latter's more restricted habitat (Appert 
1968); this habitat has been reported as being destroyed (see Threats under Long-tailed 
Ground-roller). The birds are eaten by dogs and trapped by local villagers (O. Langrand in litt. 
1984). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED A study to determine the extent and type of 
habitat destruction reported in this species's range (see under Threats) is urgently needed. A 
detailed biological study of the bird would appear likely to yield important new information in 
the realm of behavioural ecology, given its existence in groups and at least partial sex-role 
reversal. Both this and the equally remarkable Long-tailed Ground-roller, whose ranges are 
exactly coincident, merit conservation by means of a protected area. 

REMARKS This extraordinary bird occupies a monotypic genus in an endemic Madagascar 
family of little obvious affinity, both of whose other members are under threat (see relevant 
accounts). 



SLENDER-BILLED FLUFFTAIL INDETERMINATE 

Sarothrura watersi (Bartlett, 1879) Gruiformes: Rallidae 

SUMMARY This small marsh rail is known only from four well separated areas in central and 
east Madagascar, but is likely to be more widespread, and possibly more at risk from natural 
causes than from man. 

DISTRIBUTION The Slender-billed Flufftail was first described from "south-east Betsileo", 
i.e. south-central Madagascar, from which four specimens (one undated, three in December 
1875) are known (Keith et al. 1970). One of these specimens, in BMNH, is labelled 
"Fangalathova" (NJC) but this is not a locality but evidently a local name for the bird (since 
such a name is also given for the Madagascar Flufftail Sarothrura insularis in Milne Edwards 
and Grandidier 1885; see Remarks). An early map marks the south-east of "Betsileo province" 
as the region north-east and south-west of Ikongo (Locamus 1900; see also map in Deschamps 
1960). In April 1928 an immature male was collected by L. Lavauden at Analamazoatra near 
Perinet in eastern Madagascar (specimen in Grenoble: O. Langrand in litt. 1984). The species 
was subsequently found at 1,800 m near Andapa, north-east Madagascar, where 10 specimens 
were brought in by native hunters between 23 August and 7 September 1930 (Delacour 1932a, 

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Appendix 3: species accounts 

Rand 1936, Keith et al. 1970). Another two specimens are known, labelled simply 
"Madagascar" without date or name of collector (Keith et al. 1970). In 1970-1971 it was found 
in the 1,200 km 2 area around the capital, Antananarivo, central Madagascar, at three sites at 
least, and was suspected of breeding in all Cyperus marshes in this area, which is all above 
1,250 m (Salvan 1972a); however, a search around Antananarivo in the mid-1970s by three 
ornithologists (A. D. Forbes-Watson, G. S. Keith and D. A. Turner) wholly failed to rediscover 
this species, raising doubts about the validity of the records from this area (D. A. Turner in litt. 
1983). It has been speculated that this species may replace the common Madagascar Flufftail at 
higher altitudes and that it could occur on the Itremo massif (Benson et al. 1976); also that 
temperature may control its montane distribution (Rand 1936). However, Ikongo and its 
surrounding area appears to be or have been on the upper edge of the eastern rainforest belt 
and if the species was indeed collected there, and if the records from Antananarivo are in fact 
mistaken, there is a strong possibility that its distribution is determined by the distribution of 
rainforest in Madagascar. 

POPULATION Numbers are unknown. On the basis of uncorroborated observations (see 
above), density has been estimated at one pair per 2 ha of marsh, and the species perhaps 
breeds in small numbers around Antananarivo (Salvan 1972a). If these records are invalid, 
however, it is to be noted that the species has not been seen in the wild for over 50 years. 

ECOLOGY This rail inhabits small swamps (an association with Cyperus is indicated) and 
adjacent grassy areas, keeping to dense vegetation though occasionally flying short distances 
(Delacour 1932a, Rand 1936, Salvan 1972a). Food is unrecorded (Keith et al. 1970). A male 
and female in breeding condition, Andapa, September, suggest the species may be a rainy 
season breeder at that locality (Rand 1936, Keith et al. 1970). An adult with a juvenile was 
reported near Antananarivo, May (Salvan 1972a). There is no evidence of migration (Keith et 
al. 1970). 

THREATS Prior to its (uncorroborated) discovery around Antananarivo, this species was 
considered rare (Delacour 1932a) and possibly "a relict on its way to early extinction" (Keith et 
al. 1970). Antananarivo being in the most densely populated and disturbed part of Madagascar 
(Salvan 1972a), the bird may prove to be more resilient than suspected. The Laniera marshes, 
where the species has apparently bred (record of adult with juvenile, above), have been turned 
into rice-fields, and this is implied to be an ornithological disaster (Salvan 1972a); but it is not 
clear if the breeding record was made before or after this development. 

CONSERVATION MEASURES TAKEN The species has been recorded from the area now 
established as the Perinet-Analamazaotra Special Reserve, which covers 810 ha 
(Andriamampianina 1981); birds might occur in the 60,150 ha Marojejy Reserve (R.N.I, no. 12) 
(see Andriamampianina 1981), since it lies immediately north of Andapa. 

CONSERVATION MEASURES PROPOSED A detailed survey of marshes near Antananarivo 
is needed to establish whether this species is present, and at what densities; protection of 
selected sites might then be given. Searches also need to be made in the three other areas 
where birds have been found. 

REMARKS This is the least typical member of the genus Sarothrura, evidently owing to long 
isolation in Madagascar, and a genus of its own, Lemurolimnas, has been proposed, though 
regarded as unnecessary (Keith et al. 1970). Failure to confirm its presence around 
Antananarivo need not totally invalidate records from this area, since the species possibly 
shows a volatility of site-usage akin to that shown by the White-winged Flufftail S. ayresi (see 
relevant account). Although given as quoted under Distribution, the native name of this 
species is correctly "fangalatrovy" (= "stealer of yams") (J. T. Hardyman in litt. 1984). 



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SAKALAVA RAIL INSUFFICIENTLY KNOWN 

Amauromis olivieri (G. Grandidier and Berlioz, 1929) Gruiformes: Rallidae 

SUMMARY This marsh-dwelling rail is known from only three widely separated areas in the 
Sakalava country of western Madagascar, and is generally regarded as rare and localised. 

DISTRIBUTION The Sakalava Rail was first described from a single specimen (apparently 
undated) from Antsalova, west Madagascar (Grandidier and Berlioz 1929), i.e. at about 18°40'S 
44°37'E, (contra "18°28'S 44°45'E" in Benson and Wagstaffe 1972). In recent years the species 
has been seen again in the region of Lakes Masama and Bemamba by G. Randrianasolo but a 
later search of these lakes was unsuccessful (O. Langrand in litt. 1984). Soon after its first 
discovery in this region the species was found c. 300 km to the north-east at Ambararatabe 
near Soalala, roughly 16°19'S 46°04'E, where seven specimens were collected in March 1931, 
six of them along the Tsiribahina (Tsiribehino) River (Rand 1936, Benson and Wagstaffe 
1972). The only subsequent record is of a female taken from a nest at Nosy-Ambositra on the 
Mangoky River, 21°55'S 44°00'E, some 360 km to the south of the type-locality, on 9 March 
1962 (Benson and Wagstaffe 1972). This record has done nothing to modify the description of 
the species, over 50 years ago, as strictly localised (Delacour 1932a), which clearly implied that 
it had been looked for in other areas and found absent. From its behaviour (see Ecology 
below), it would seem less easy to overlook than, e.g., the Slender-billed Flufftail Sarothrura 
watersi (see relevant account), and new localities for it may prove to be few. However, large 
areas of apparently suitable but inaccessible habitat do exist (D. A. Turner pers. comm. 1983). 

POPULATION Numbers are unknown. 

ECOLOGY Birds along the Tsiribahina River at Ambararatabe were found standing on or 
running over floating vegetation on a narrow, deep stream bordered with tall coarse grass 
locally called "bararata" (apparently the reed Phragmites communis: see Benson and Wagstaffe 
1972); though not very shy or active, the birds kept close to the "bararata" and retreated there 
for shelter (Rand 1936). A bird was also seen on a floating log in a flooded valley clearing; on 
26 March a male and female were seen with two well-grown young (Rand 1936). The nest at 
Nosy-Ambositra was some 50 cm above ground level in bulrushes Typha angustifolia near 
water, in a marshy area with stretches of open water, with bulrushes, water-lilies Nymphaea 
stellata and Phragmites communis dominant (Benson and Wagstaffe 1972). The nest held two 
eggs, probably a complete clutch (Benson and Wagstaffe 1972). 

THREATS The species's very restricted distribution, as currently known, exposes it to a 
variety of potential threats. The eggs of the only recorded nest were eaten by local people 
(Benson and Wagstaffe 1972) and it is possible that populations could suffer locally from 
systematic exploitation for food. Rice-growing is reportedly beginning to alter Lake Bemamba 
(O. Langrand in litt. 1984). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED A faunal reserve has been urged for the 
Antsalova region, this species's type-locality; for information on this proposal and on the 
possible importance of wetlands around Cap St. Andre, see Conservation Measures Proposed 
under Madagascar Teal Anas bernieri. Research on the Teal's distribution could incorporate 
fieldwork to locate populations of this rail and to determine the threats it may face. 

REMARKS Although commonly placed in the genus Porzana, the Sakalava Rail has been 
found to show close affinity to the African Black Crake Amauromis (Limnocorax) flavirostris 
(Benson and Wagstaffe 1972). 



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Appendix 3: species accounts 
MADAGASCAR PLOVER RARE 

Charadrius thoracicus (Richmond, 1896) Charadriiformes: Charadriidae 

SUMMARY This shorebird is apparently restricted to coastal grassy areas of south-west 
Madagascar where it is greatly outnumbered (and possibly outcompeted) by Kittlitz's 
Plover Charadrius pecuarius. 

DISTRIBUTION The Madagascar Plover is now (largely or exclusively) confined to coastal 
south-west Madagascar. It was, however, first described from Loholoka (21°44'S 48°12'E) and 
the Fanantara estuary (20°51'S 48°28'E) on the east coast of Madagascar (i.e. between 
Mahanoro and Manakara), when other specimens from the south-east coast were also 
mentioned (Richmond 1896,1897; coordinates in Office of Geography 1955); there is also a 
specimen collected by A. Lantz and received by MNHN in 1882 labelled as from the south-east 
coast (NJC). In the present century it has only been reported with certainty - other than an 
anomalous inland record of four 60 km from Antananarivo in January 1971 (Salvan 1971) - 
from the south-west coast between Morondava and Androka. The species has recently been 
reported without comment from Morondava (O. Langrand in litt. 1984), though this is much 
the most northerly coastal record, birds not otherwise being known to extend beyond the 
Maintapaka estuary (north of the Mangoky River) (Appert 1971a). Thirteen sites were mapped 
for the species in the Morombe/Mangoky delta area in the 1960s (Appert 1971a) and several 
more were found between Morombe and Lake Tsimanampetsotsa, July/ August 1972 (Dhondt 
1975). Previous records are from Toliara airstrip (Milon 1950), Lake Tsimanampetsotsa (Bangs 
1918, Milon 1950), "Nosy Asatra to Beheloka" (Bangs 1918; see Remarks), Androka (Uinta 
estuary) (Delacour 1932a, Rand 1936) and Nosy Mborono (Nosimborona), off Androka (Milon 
1948). The species is not found at Lake Ihotry or near Antsalova (Dhondt 1975). 

POPULATION There are no estimates, but on published evidence the total number must be 
low, possibly under a thousand. The largest flocks reported are of 33 (Appert 1971a) and 16 
(Dhondt 1975); the relatively few other records are all in (usually low) single figures, e.g. only 
three were found by the Mission Franco- Anglo- Americaine, 1929-1931, after which the 
species was judged very rare (Delacour 1932a). A two-day survey of the area between 
Morombe and Befandefa, July 1972, recorded a total of seven birds at four sites; 76 Kittlitz's 
Plover Charadrius pecuarius were found in the same places (Dhondt 1975). A two-day survey 
at Lake Tsimanampetsotsa, August 1972, recorded a total of 39 Madagascar Plovers at eight 
sites; at one of these, where several hundred plovers were probably present, 37 were pecuarius 
and only seven thoracicus (Dhondt 1975). The species is, however, reported to be "rather 
common" in the Morombe region (O. Langrand in litt. 1984). 

ECOLOGY It inhabits short-grass areas near the coast, also flat margins of saltwater expanses 
and pools, occurring less often on sand- or mudflats (Appert 1971a). At least in July and 
August, it appears to prefer drier areas than Kittlitz's Plover and even to avoid flooded 
grassland (Dhondt 1975). The stomach of a specimen in MNHN held large and small insects 
("not grasshoppers") (NJC). Eggs have been recorded in November (Appert 1971a) and January 
(Milon 1950), young in December (Appert 1971a) and August (Bangs 1918, Milon 1950). 
Breeding and general biology is evidently close to Kittlitz's Plover (see Appert 1971a, Keith 
1980; for Kittlitz's, see Cramp and Simmons 1983). 

THREATS The reasons for this species's rarity are unclear. What is certain is that Kittlitz's 
Plover is more recent in Madagascar (Keith 1980), and its relative numerical superiority and 
much wider distribution suggest that it may compete successfully with thoracicus. 
Hybridisation has not been recorded. 

CONSERVATION MEASURES TAKEN The species occurs in the Lake Tsimanampetsotsa 
Nature Reserve (R.N.I, no. 10) (Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED A study of the status, distribution and 
year-round ecological requirements of this species is clearly essential in order to determine the 
measures needed for its survival. Any such study should include a survey of the east coast of 
Madagascar from Toamasina southwards. For the need for a general ornithological survey of 

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An environmental profile of Madagascar 

this coast and its wetlands, see under Conservation Measures Proposed for Madagascar 
Teal Anas bernieri. 

REMARKS Birds with black breast-bands were not recognised as a species distinct from 
Kittlitz's Plover until 1896 (see Richmond 1896), and it is possible that museum collections 
hold specimens of thoracicus from before that date whose locality data would be of value in 
determining the extent of its (at least former) distribution. Neither "Nosy Asatra" nor 
"Beheloka" are gazetteered (in Office of Geography 1955), but it is clear (from Agassiz 1918 
and Bangs 1918) that they must lie between Toliara and Lake Tsimanampetsotsa, along the 
coast, and indeed a Nosy Satrana and Pointe de Beheloka are marked in this stretch of coastline 
(in IGNT 1964), at 23°43'S 43°38'E and 23°55'S 43°40'E respectively. 



SNAIL-EATING COUA EXTINCT 

Coua delalandei (Temminck, 1827) Cuculiformes: Cuculidae 

SUMMARY This large terrestrial cuckoo is the only bird (other than the elephantbirds 
Aepyornithidae) in Madagascar generally believed to have become extinct. There is a very 
remote possibility that it survives. 

DISTRIBUTION The Snail-eating Coua is known chiefly from Nosy Borah (He de 
Sainte-Marie) off the northern east coast of Madagascar (Sganzin 1840, Ackerman 1841). The 
species is also repeatedly stated to have occurred on the mainland opposite Nosy Borah, 
especially on the immediately adjacent Pointe-a-Larree (Milne Edwards and Grandidier 1879, 
Milon el al. 1973; also Hartlaub 1877, Delacour 1932a, Rand 1936) and, perhaps owing to its 
reported survival in the deepest forests of the region between Fito and Maroantsetra (Lavauden 
1932), its mainland range has been guessed as "from the head of Antongil Bay southward to 
Tamatave (=Toamasina)" (Peters 1940). However, it has been pointed out that "there are no 
exact records of the provenance of mainland specimens" (Greenway 1967), and indeed it is 
nowhere clear that any specimen is known to have come from anywhere other than Nosy 
Borah. At least 13 specimens (two each in BMNH and MNHN, one each in MCZ, AMNH, 
ANSP, RMNH, SMNS, NHMW, Liverpool, Antananarivo and IRSNB: see Remarks) are known 
to exist (Hartlaub 1860, Delacour 1932a, Rand 1936, Greenway 1967, Benson and Schtiz 1971, 
Schifter 1973, Morgan 1975, NJC), the origin of many of which seems likely to have been 
Nosy Borah, as it is recorded that specimens from there were dispersed to various museums 
(Sganzin 1840). Nevertheless, plate 65 in Milne-Edwards and Grandidier (1876) maps the 
distribution of this species as the eastern rainforest from the latitude of Toamasina north to 
that of Nosy Borah (but not Nosy Borah itself); the authority for such a distribution is not 
given. A record of the species as a "waterbird" at Lake Alaotra (Baron 1882) is presumably in 
error. 

POPULATION The extinction of this species is probable (as judged in Delacour 1932a, Rand 
1936, Milon 1952, Greenway 1967) but not certain (contra Day 1981). None has been reported 
with certainty since 1834 (Greenway 1967), although the dates of Ackerman's three-year stay 
(see Ackerman 1841) are not clear and there are three specimens which could have been 
collected after this date, though not later than 1837, 1840 or 1850 respectively (Benson and 
Schuz 1971, Schifter 1973, Morgan 1975). The species was "not very rare" on He de 
Sainte-Marie in 1831-1832 (Sganzin 1840), which may perhaps be the source of the statement 
in 1860 that it was "not rare on the east coast" (Hartlaub 1860); however, no trace of it could be 
found during six month's exploration in 1865 and it was therefore judged very rare (Milne 
Edwards and Grandidier 1879; also Jouanin 1962). Following the failure of the Mission 
Franco-Anglo-Americaine to find it in 1929-1931, and the failure of the offer of a large 
reward to the procurer of a specimen in 1932 (see Greenway 1967), it was pronounced 
probably extinct (Delacour 1932a, Rand 1936). Nevertheless, at just this time a "very reliable 
native who knew exactly what bird was being referred to" reported that the species stili 
survived on the mainland but was very rare and very shy (Lavauden 1932). Much of the area 
in question was not visited by the Mission Franco-Anglo-Americaine, and has not apparently 

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Appendix 3: species accounts 

been searched subsequently, and it is accepted that the species might conceivably survive in a 
few remote undisturbed patches (Greenway 1967, Milon et al. 1973). Survival on Nosy Borah 
is ruled out as all the original forest has long since been cleared (Lavauden 1932, Daumet 1937, 
Petter 1963, Keith et al. 1974). 

ECOLOGY This bird is or was a ground-haunting rainforest-dweller, subsisting on molluscs 
(Sganzin 1840). An account of its method of breaking snail shells, based on observations in an 
aviary and apparently also in the wild, has been provided along with brief details of its 
behaviour and voice (Ackerman 1841). 

THREATS Habitat destruction was clearly the chief cause of its disappearance from Nosy 
Borah (Lavauden 1932, Petter 1963, Keith et al. 1974), and was identified as the chief threat to 
its existence on the mainland as long ago as 1932 (Lavauden 1932): most of the lowlands 
between Toamasina and Maroantsetra were devoid of forest at the end of the 1960s (Keith et 
al. 1974). Occasional snaring by natives was reported (Lavauden 1932) and this was 
presumably quite easy at a time when the species was more numerous, and may have played a 
part in its decline; it was reportedly hunted as much for feathers as for food (Keith et al. 
1974). Shell remains at certain localities within the forest may have betrayed the presence of 
birds to hunters (A. D. Forbes-Watson pers. comm. 1984). A recent account gives a cause of 
extinction as "introduced rodents" (Day 1981): while there appears to be no direct evidence for 
this, it is conceivable that rats critically reduced the mollusc fauna in key areas and this 
indirectly contributed to the species's disappearance. For evidence of rats in eastern forests, 
see Threats under Brown Mesite Mesitornis unicolor. 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED It needs to be properly established, through a 
reexamination of museum material and records, whether the species occurred on mainland 
Madagascar. Even if this cannot be done, the ornithological surveys that are needed for other 
reasons in the Sihanaka and other remaining forests between Fito and Maroantsetra (see 
Conservation Measures Proposed under Madagascar Serpent Eagle Eutriorchis astur) should 
certainly be weighted towards tracking down evidence of this bird's survival. 

REMARKS It is to be observed that if the White-breasted Mesite Mesitornis variegata went 
unknown from 1834 to 1929 (see relevant account) and the Yellow-bellied 
Sunbird-asity Neodrepanis hypoxantha from 1929 almost to the present day (but is not extinct: 
see relevant account), it is certainly conceivable - if less likely - that the Snail-eating Coua 
could have survived undetected over the same 150-year period. The specimen in IRSNB, 
whose existence has not previously been announced, was acquired by the museum in 1839 and 
according to the catalogue it was captured or collected in "Madagascar" in 1832 (P. Devillers 
pers. comm. 1983), this date perhaps rendering it likely to have come from Sganzin on Nosy 
Borah (see Distribution). 



MADAGASCAR RED OWL INDETERMINATE 

Tyto soumagnei (Milne Edwards, 1878) Strigiformes: Tytonidae 

SUMMARY This owl is known with certainty from rainforest only in eastern central 
Madagascar, and has been seen only once in the past 50 years. 

DISTRIBUTION The Madagascar Red Owl inhabits the eastern region of Madagascar in the 
circle whose diameter runs between Toamasina and Antananarivo. It does not occur 
"throughout Madagascar" (contra Burton 1973). The type-specimen was collected in 1876 on 
the east coast near Toamasina (Milne Edwards and Grandidier 1879) and a specimen from 
around Antananarivo (no date) came to the British Museum in 1879 (Sharpe 1879); as this 
specimen is catalogued as being collected by "Lorimer" (NJC) it presumably cannot be the bird 
sent back, also in 1879, by Humblot but which is not listed as going to MNHN (Humblot 1882) 

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and indeed cannot be found there (NJC,SNS). There are two other nineteenth century 
specimens (in BMHN), one from "the upper forest of Eastern Imerina" in March 1893, one 
from "Merimitatra" (the label also states "between the two forests"), east Madagascar, January 
1895 (Wills 1893, NJC); the former area has been cleared of forest (D. A. Turner pers. comm. 
1983), but a place bearing the latter name is marked (in Locamus 1900) as a comparatively 
large settlement (now abandoned or re-named: not in Office of Geography 1955 or on recent 
maps) east of Anjozorobe, at roughly 18°25'S 48°05'E, on the upper western slopes of the 
Mangoro valley and thus between the two belts of forest bordering the valley (Sihanaka forest 
in the east, Angavo escarpment forest in the west). One collector obtained only three birds of 
this species in 40 years on Madagascar (two of these specimens were destroyed in 1927) 
(Lavauden 1932); all three were found in Sihanaka forest (Delacour 1932a). Two specimens (a 
pair) were shot in March 1930 at Analamazoatra, near Perinet (Lavauden 1932), and another 
was taken near Fito, Sihanaka forest, on 15 February 1934 (Allen and Greenway 1935). The 
only subsequent record has been of a bird in deep mountainous rainforest (1,200-1,800 m) a 
day's walk from the nearest motorable road, Fierenana district (c. 65 km north of Perinet), in 
1973 (King 1978-1979, J. I. Pollock in litt. 1983). The species is also reported as occurring on 
the Masoala peninsula (Milon et al. 1973) but evidence for this - although it seems likely - has 
not been traced. 

POPULATION Numbers are unknown, but the species has always appeared to be extremely 
rare (e.g. Delacour 1932a, Lavauden 1932, Milon et al. 1973). 

ECOLOGY This owl inhabits humid rainforest and is strictly nocturnal, reportedly living in 
isolated pairs and feeding on frogs caught in clearings (Lavauden 1932). There are no other 
data, but it is to be observed that at least three specimens have come from localities 
(Toamasina, Antananarivo and Merimitatra) apparently outside heavily forested areas; however, 
it is not known to occur in grassland (contra Burton 1973). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). 

CONSERVATION MEASURES TAKEN The species has been recorded from the area now 
established as the Perinet- Analamazoatra Special Reserve, which covers 810 ha 
(Andriamampianina 1981). The Madagascar Red Owl is listed on Appendix I of CITES, to 
which Madagascar is a party. 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. 

REMARKS Although originally placed in its own genus (Heliodius), this species is clearly a 
small, dark reddish-orange barn owl Tyto (Lavauden 1932, Allen and Greenway 1935; also 
Sharpe 1879). 



SHORT-LEGGED GROUND-ROLLER RARE 

Brachypteracias leptosomus (Lesson, 1833) Coraciiformes: Brachypteraciidae 

SUMMARY This roller is confined to deep rainforest in the centre and north-east of 
Madagascar, and is widely considered rare. It is threatened by forest destruction. 



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DISTRIBUTION On current limited knowledge, the Short-legged Ground-roller occurs in two 
discrete general areas of Madagascar, in the north-east (Marojejy to around Maroantsetra) and 
central-east (chiefly Sihanaka forest). Records from the north-east are from the Marojejy 
Nature Reserve, September 1972 (Benson et al. 1976), around Antanombo Manandriana, one 
day's march west of Andapa, 1930 (Delacour 1932a, Rand 1932,1936), around 
"Ambolumarahavany" (see Remarks), two days' march north-east of Maroantsetra, 1930 
(Delacour 1932a, Rand 1932,1936), and around Bevato, 40 km north-west of Maroantsetra, 
1930 (Delacour 1932a, Rand 1932,1936), Maroantsetra (specimen in SMF: NJC), Mananara, 
November 1876, and Savary (for location of which see Distribution under Brown 
Mesite Mesitornis unicolor), November 1877 to April 1878 (specimens in RMNH: NJC), and 
from the Masoala peninsula (Turner 1984). In the central-east, the species is known from 
Sihanaka forest (Delacour 1932a), Perinet, August 1982, at 950 m (O. Langrand in lilt. 1984), 
the Toamasina region, September 1913 (specimen in SMF: NJC), Fanovana (Delacour 1932a) 
and the east Imerina forest (Wills 1893, Oberholser 1900), these last two areas having now been 
cleared (D. A. Turner pers. comm. 1983) as presumably has that around Toamasina. There is 
also a record from near Ampasimbe (Newton 1863; see Remarks), and a skin in BMNH, 
undated and labelled "Sambririna": no such locality can be traced (in Office of Geography 
1955), but the possibility that "Sambirano" is intended - i.e. the area of humid forest in the far 
north-west - cannot be ignored. Two other localities, "Ambore" and "Ankoraka 
Sahambendrana" (specimens in ZFMK and MNHN respectively: NJC) cannot be traced. 

POPULATION This species has been considered commoner than the largely sympatric Scaly 
Ground-roller Brachypteracias squamiger (see relevant account), and over two years 42 
specimens were collected or acquired as against 20 of the latter (Delacour 1932a,b). 
Nevertheless the Short-legged Ground-roller has a somewhat more restricted range and within 
this has consistently been regarded as rare in some degree (Hartlaub 1877, Milne Edwards and 
Grandidier 1881, Rand 1936, Lavauden 1937, Milon el al. 1973). A recent study has suggested 
that all ground-rollers have been thought rarer than they are, since their silence and secretive 
behaviour lead them to be "completely overlooked"; this species is considered "shy though not 
uncommon" (Turner 1984). 

ECOLOGY It inhabits heavy rainforest, "frequenting low, wet places where the trees cast a 
continual shade and the ground-cover of spindly saplings leaves the damp forest floor nearly 
bare" (Rand 1936). Although it is considered terrestrial, one observation was of a bird that 
perched on horizontal strands of vines and in small trees, remaining immobile for minutes on 
end, with short fast flights between perches (Benson et al. 1976), another was of a bird which, 
when flushed from the ground, flew up to a tree and hid behind branches (Dresser 1893), 
while a recent study suggests it is in fact much the most arboreal of the ground-rollers (Turner 
1984). It has been reported to scratch at moss and dead leaves with its feet like a gallinaceous 
bird, to uncover beetles, ants, larvae, millipedes, pill-millipedes, ant-lions, worms and small 
reptiles (Milne Edwards and Grandidier 1881; hence Milon et al. 1973). Of eight stomachs, one 
held a snake; two, chameleons; one, beetles; two, caterpillars; four, other insects; one, a snail 
(Rand 1936). Two other stomachs held large ants plus beetle remains (Milon et al. 1973) and 
tenebrionid beetles (Benson et al. 1976) respectively. Natives reported it to be a night-feeding 
bird (Sharpe 1871) and it is said to be at least partly nocturnal (Hildebrandt 1881) and locatable 
in the early morning and evening (Milne Edwards and Grandidier 1881). It is solitary except 
in breeding season, when it occurs in pairs (Milne Edwards and Grandidier 1881). Birds breed 
in December, excavating the nest in a tunnel (c. 1 m) in a bank (Milon et al. 1973); they are 
also reported to nest in holes in trees (Dresser 1893). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). 

CONSERVATION MEASURES TAKEN The species occurs in the Marojejy Nature Reserve 
(R.N.I, no. 12), which covers 60,150 ha, and in the Perinet- Analamazaotra Special Reserve, 
which covers 810 ha (Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED Full protection for all ground-rollers has been 
called for (Salvan 1970). Immediate and effective protection of as much remaining rainforest 

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as possible would almost certainly guarantee the survival of this and all other 
rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. This species requires study to determine the basic 
aspects of its ecology and whether or not it is migratory: in view of native reports that it 
hibernates (Dresser 1893), it seems likely that some movement takes place, and an 
understanding of this may be crucial to its long-term conservation. 

REMARKS From context, Ampasimbe is evidently the locality on the main road from 
Antananarivo to Andevoranto, at 18°58'S 48°40'E, well outside the main rainforest block (see 
Office of Geography 1955, IGNT 1964); however, an earlier map indicates that a small belt of 
forest, named Madilo, crossed the road near Ampasimbe (see Locamus 1900), though this is 
presumably now all cleared. "Ambolumarahavary" is presumably identical to 

"Ambohimarahavary" (see Distribution under Madagascar Serpent Eagle) and is probably, 
correctly, "Ambolomirahavavy" (J. T. Hardyman in lilt. 1984), though in fact none of these 
names can be traced. 



SCALY GROUND-ROLLER RARE 

Brachypteracias squamiger Lafresnaye, 1838 Coraciiformes: Brachypteraciidae 

SUMMARY This roller is confined to deep rainforest in the centre and north-east of 
Madagascar, and is widely considered rare. It is threatened by forest destruction, by village 
dogs and by human exploitation for food. 

DISTRIBUTION The Scaly Ground-roller occurs throughout the eastern rainforests of 
Madagascar. Records are from (north to south) Marojejy (Benson et al. 1976), Andapa (Rand 
1936), around Maroantsetra (Rand 1936, O. Langrand in litt. 1984; see Remarks), 
"Mointenbato" (Fisher 1981), i.e. Maintimbato, and Savary (for location of both see Distribution 
under Brown Mesite Mesitomis unicolor), December 1877 to April 1878 (specimens in RMNH 
and MNHN: NJC), the Masoala peninsula (B.-U. Meyburg pers. comm. 1983, Turner 1984), the 
Soamianina (= "Semiang", "Tsimianona") River (type-locality, opposite Nosy Borah) (Hartlaub 
1877, Milne Edwards and Grandidier 1881), Sihanaka forest (Delacour 1932a, Rand 1936), 
Perinet (Webb 1954), Analamazoatra (specimen in Grenoble: O. Langrand in litt. 1984), Rogez 
(Benson et al. 1976), the Toamasina region, July 1912 and October 1913 (specimens in MNHN 
and SMF: NJC) and south-east Madagascar (Dresser 1893). Its occurrence in the south-east has 
been entirely overlooked this century, but was substantiated by four specimens (see Dresser 
1893). One specimen in BMNH is labelled "Voolaly, S. E. Madagascar" (untraceable) and dated 
February 1872 (NJC); another in SMF was collected on 8 October 1931 at Eminiminy, 
south-east Madagascar (NJC), gazetteered as at 24°41'S 46°48'E (Office of Geography 1955) 
and mapped as on the eastern boundary of the Andohahela Nature Reserve (in IGNT 1964). 
Of two specimens in BMNH collected by J. Audebert and dated February 1879, one has an 
illegible label (see Remarks), while the other is from "Antsondririna" (NJC): the only 
gazetteered locality of this name is in the far north-east, at 13°00'S 49°41'E (Office of 
Geography 1955), and thus much the most northerly record (if correct) for the species. 

POPULATION This species has been widely considered rare in some degree: "very rare" 
(Sganzin 1840), "quite rare" (Milne Edwards and Grandidier 1881), "everywhere rare" (Rand 
1936; also Delacour 1932a). Nonetheless, a recent study has suggested that all ground-rollers 
have been thought rarer than they are, since their silence and secretive behaviour leads them to 
be "completely overlooked" (Turner 1984). The species was seen almost daily on the Masoala 
peninsula in October 1980 (B.-U. Meyburg pers. comm. 1983). 

ECOLOGY The Scaly Ground-roller is a ground-adapted bird of heavy, deep-shaded 
rainforest with sparse undergrowth (Rand 1936; also Hartlaub 1877, Benson et al. 1976), 

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Appendix 3: species accounts 

considered the terrestrial counterpart of the somewhat arboreal Short-legged 
Ground-roller Brachypteracias leptosomus (see relevant account); when disturbed, it either flies 
a few yards or runs a few steps, then stands quietly watching the intruder (Turner 1984). 
Native reports that it is nocturnal (Sharpe 1871) have not been proven; and though several 
ground-rollers appear to be most active at dusk (Turner 1984), recent observations on this 
species suggested it to be active throughout the day (O. Langrand in lilt. 1984). Of five 
stomachs, four contained large terrestrial insects, one a spider (Rand 1936); another held ants 
and scarabaeid beetles (Benson et al. 1976). Prey seen taken includes ground-beetles, ants, 
caterpillars, centipedes, earthworms, snails, small frogs; Lepidoptera and Diptera are also 
hawked in flight; whether or not the Short-legged Ground-roller scrapes at the leaf-litter with 
its feet to uncover its prey, as reported (see relevant account), the Scaly Ground-roller only 
ever uses its bill for such purposes (O. Langrand in lilt. 1984). Breeding evidently occurs in 
September (see Benson et al. 1976); a nest-hole probably of this species consisted of a tunnel 
less than a metre long, with a chamber lined with dead leaves and earthy pellets, built into a 
bare, sloping bank in deep forest (Benson et al. 1976). A nest with young was found on 4 
November 1982, 50 km north-west of Maroantsetra at 350 m (O. Langrand in lilt. 1984). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). The species is also threatened by young villagers, who trap 
birds and catch them in the nest, and by village dogs which also catch birds (O. Langrand in 
lilt. 1984). 

CONSERVATION MEASURES TAKEN The species occurs in the Marojejy Nature Reserve 
(R.N.I, no. 12), which covers 60,150 ha, and presumably in the Perinet-Analamazaotra Special 
Reserve, which covers 810 ha, and the Andohahela Nature Reserve (R.N.I, no. 11), which 
covers 76,020 ha (Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED Full protection for all ground-rollers has been 
called for (Salvan 1970). Immediate and effective protection of as much remaining rainforest 
as possible would almost certainly guarantee the survival of this and all other 
rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. 

REMARKS For exact localities where the species has been found around Maroantsetra, see 
Rand (1932). The illegible locality on the Audebert label (see Distribution) is possibly 
"Ampirina", but no such name has been traced (on Locamus 1900 or in Office of Geography 
1955), although it must be fairly close to Antsondririna to have been visited in the same month 
in 1879. 



RUFOUS-HEADED GROUND-ROLLER RARE 

Atelornis crossleyi Sharpe, 1875 Coraciiformes: Brachypteraciidae 

SUMMARY This roller is confined to deep rainforest in the centre and north-east of 
Madagascar, and is widely considered rare. It is threatened by forest destruction. 

DISTRIBUTION On current limited knowledge, the Rufous-headed Ground-roller occurs in 
two discrete general areas of Madagascar, in the north-east (Tsaratanana massif, Marojejy 
Reserve, and Andapa) and central-east (in a circle whose diameter runs from Antananarivo to 
Toamasina); it has also been recorded in two more southerly general forest areas in 
south-central Madagascar. Records from the north-east appear to be based on only three 
specimens, one from Tsaratanana massif at 1,500 m in 1966 (Albignac 1970), one from 
Ambodifiakarana, in the Marojejy Nature Reserve, at 1,600 m in 1958 (Griveaud 1960b, 

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An environmental profile of Madagascar 

Benson et al. 1976), and one from Antanombo Manandriana, one day's march west of Andapa, 
at around 1,800 m in 1930 (Delacour 1932a, Rand 1932,1936). The bird is next recorded some 
400 km to the south from the Sihanaka forest (Delacour 1932a, Rand 1936), including Didy 
(Milon et al. 1973) and Fito (Benson et al. 1976; specimens in MNHN: NJC). Birds have also 
been seen, collected or acquired from natives in "Foret Ruanaka" (untraceable: possibly a 
mistake for "Sianaka") in the Brickaville district, Vohibazaha forest and Lakato forest 
(specimens in MR AC: NJC), Analamazaotra, and near Perinet (Lavauden 1932, O. Langrand in 
litt. 1984, A. D. Forbes-Watson pers. comm. 1984; see Conservation Measures Taken), 
Fanovana (Delacour 1932a) and the east Imerina forest (Dresser 1893, Rothschild 1895, 
Oberholser 1900), though this and the forest at Fanovana are now cleared (D. A. Turner pers. 
comm. 1983). The type-locality, first reported as "Ampasmonhavo" (Sharpe 1875), then 
"Ampasimanavy" (Milne Edwards and Grandidier 1881), is almost certainly therefore 
Ampasimaneva, on the Mangoro River, at 19°24'S 48'04'E (see Remarks under Madagascar 
Serpent Eagle Eutriorchis astur). The most southerly records are from south-central 
Madagascar in "the forest land that lies between the Betsileo and Tanala.. .[which] covers the 
eastern side of the mountains along the edge of the central plateau. ..[and] is thick and dense, 
about fifteen or twenty miles in width" (Deans Cowan 1882) and, in 1984, from the Vondrozo 
region (O. Langrand pers. comm. 1984). 

POPULATION The species is widely considered rare in some degree (Richmond 1897, 
Lavauden 1932, Rand 1936, Griveaud 1960b) but, from the number of skins in one local 
collection around 1930, it was evidently then "not uncommon" in Sihanaka forest (Rand 1936; 
also Delacour 1932a) and indeed "not rare" at Didy, presumably around 1970 (Milon et al. 
1973). It was listed as common in the Betsileo/Tanala border forest a century ago (Deans 
Cowan 1882). A recent study has suggested that all ground-rollers have been thought rarer 
than they are, since their silence and secretive behaviour lead them to be "completely 
overlooked"; however this is considered the rarest and least known species (Turner 1984), a 
view endorsed by recent observations (O. Langrand in litt. 1984). 

ECOLOGY "This bird probably frequents the ground in the heavy forest; one stomach 
examined contained insect remains" (Rand 1936). It is reported to nest in holes in the ground 
(Dresser 1893). A not fully grown bird collected in late March (specimen in MNHN: NJC) 
suggests a December/January breeding season. In central-east Madagascar, the species 
seemingly disappears during the winter months (May/August) (Dresser 1893), though there are 
in fact specimens (in BMNH, MNHN and SMNS) from Sihanaka taken in May and August 
(NJC). There appears to be no other information specifically on this bird, although it has been 
listed as characteristic of secondary forest dominated by Ravenala madagascariensis, such 
forest mostly occurring from sea-level to 500 m (Lavauden 1937); the basis and validity of this 
assertion are unknown. 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle). Two areas of forest where it was known to occur have been felled (see 
Distribution). 

CONSERVATION MEASURES TAKEN The species evidently occurs in both the Marojejy 
Nature Reserve (R.N.I. no. 12), which covers 60,150 ha, and the Tsaratanana Nature Reserve 
(R.N.I, no. 4), which covers 48,622 ha (Andriamampianina 1981), although its status in both 
(one record each) appears precarious. Observation of this ground-roller several kilometres 
from and c. 100 m higher than the Perinet- Analamazaotra Special Reserve has been taken to 
suggest that the reserve may be too low to be of great value in helping to preserve this species 
(A. D. Forbes-Watson pers. comm. 1984). 

CONSERVATION MEASURES PROPOSED Full protection for all ground-rollers has been 
called for (Sal van 1970). Immediate and effective protection of as much remaining rainforest 
as possible would almost certainly guarantee the survival of this and all other 
rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 

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Appendix 3: species accounts 

be extended to include searches to locate it. This species requires study to determine the basic 
aspects of its ecology, particularly in relation to its apparent migrations (see Ecology above), an 
understanding of which is probably crucial to its long-term survival. 



LONG-TAILED GROUND-ROLLER RARE 

Uratelornis chimaera Rothschild, 1895 Coraciiformes: Brachypteraciidae 

SUMMARY This remarkable terrestrial bird of restricted range within the subdesert region of 
south-west Madagascar, although numerically safe at present, appears to enjoy no protection 
whatever. 

DISTRIBUTION The Long-tailed Ground-roller is restricted to a coastal strip between the 
Mangoky and Fiherenana rivers, south-west Madagascar, ranging up to 80 m altitude (Appert 
1968, Milon et al. 1973). Most sites known for the species within this area up to 1966 have 
been listed (see Appert 1968, but also Oustalet 1899, Menegaux 1907, Bangs 1918), with the 
conclusion that its distribution coincides with that of Didierea woodland (Appert 1968). It 
therefore has stricter habitat requirements and a lower altitudinal tolerance than the closely 
sympatric Subdesert Mesite Monias benschi (see relevant account). If the altitudinal limit 
quoted above is accurate, even only roughly, it cannot occur much more than 30 km inland in 
the southern half of its range (see 100 m contour in Army Map Service 1968), while in the 
northern half it has not been recorded east of Lake Ihotry (Appert 1968) other than at Mamono 
village near Ankida (O. Langrand in litt. 1984). Moreover, the revelation that some seasonal 
movement may occur (see Ecology below) suggests that birds range beyond the currently 
known limits or that they occupy only parts of their known range at any given season. 
Evidence of its presence beyond the confines of the Mangoky and Fiherenana, e.g. on 
Montagne de la Table south of Toliara (Rand 1932; see Remarks), is lacking, and the report 
that it occurs south to Cap Sainte-Marie (Lavauden 1937) is in error. 

POPULATION In the south between Toliara and Manombo, at the turn of the century, it was 
found in good numbers mainly at Ambolisatra (Menegaux 1907), and this area is obviously still 
important (see Appert 1968). It was found to be fairly common around Lake Ihotry in 1929 
(Rand 1932,1936) but was apparently becoming rare there in the 1950s (Griveaud 1960a) and 
was judged to be "extremely rare" from second-hand information in the early 1960s (Petter 
1963). In 1968 it was described as "one of the rarest birds in the world" (Appert 1968). A 
survey (presumably around 1970) concluded that "the total population between Tulear 
(=Toliara) and Lake Ihotry is not more than 500 pairs, and nearer 250 pairs with an 80% 
probability" (Milon et al. 1973). More recently, a repeated visitor to its area of distribution has 
suggested that "in areas of undisturbed habitat it is common, and may even be termed locally 
abundant, particularly in the area of dense Didierea woodland some 30 km north of Tulear" 
(Turner 1984), a judgement supported by another recent observer (O. Langrand in litt. 1984). 

ECOLOGY The Long-tailed Ground-roller inhabits very arid areas in low, generally fairly 
dense deciduous woodland, always with (mostly herb- and grass-free) sandy soil (a prerequisite 
for nesting); it is commonly found in association with the cactus-like Didierea 
madagascariensis and the sporadic Euphorbia stenoclada, although absent from Didiereacovered 
dunes, which are probably too loose and too little shaded (Appert 1968). It feeds almost 
exclusively on terrestrial invertebrates (e.g. beetles, grasshoppers, cockroaches, woodlice, 
caterpillars, ants), typically by rummaging in leaf-litter beneath a bush or tree (Appert 1968; 
also Oustalet 1899, Rand 1936, Milon et al. 1973, and specimen-labels in MNHN: NJC); a 
low-flying butterfly was also once seen taken (Appert 1968). Birds are active (singing and 
feeding) at night, at least at times (Appert 1968; also Turner 1984); singing occurs commonly in 
late winter (August/September) (O. Langrand in litt. 1984). The species keeps largely to the 
ground, running powerfully and flying rarely, but typically calls from a low (up to 3 m high) 
horizontal perch (Appert 1968). Although it is stated to occur in small family groups 
(Lavauden 1937; hence Petter 1963, Milon et al. 1973), this can only happen for a short period 
in the year, as the species is otherwise reported to occur singly over the southern winter, and 

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always in pairs from the start of the breeding season (October or earlier through to January) 
(Appert 1968; also Rand 1936). In one area where it was studied (30 km north of Toliara), 
birds appeared to be seasonal in occurrence, being present from September to April but 
generally absent from May to August (Turner 1984). The nest-hole is excavated by both birds 
in flat or slightly sloping ground, with a tunnel extending to up to 120 cm (Appert 1968); a 
report of nesting in steep river-banks (Lavauden 1937) has been doubted (Appert 1968; also 
O. Langrand in lilt. 1984). Clutches reputedly consist of three or four eggs (Lavauden 1937), 
this being supported by a record of three juveniles evidently from one brood (Rand 1936). 

THREATS The restricted range of this species must be a source of permanent concern and 
vigilance for its welfare; such concern is compounded by its apparently migratory behaviour 
(see Ecology above), which doubles the risk it faces from any habitat destruction. This species 
was hunted by herdsmen with blowpipes at the turn of the century (Menegaux 1907) and 
natives were trapping birds and digging out nests in the 1950s and 1960s (Griveaud 1960a, 
Appert 1968); trapping by local villagers is still being practised (O. Langrand in litt. 1984). 
Some 20 years ago, the species's habitat was reported to be in a "critical situation ... being more 
and more broken up and ... in process of rapid extinction" (Petter 1963). Subsequent observers 
(Appert 1968, Turner 1984) have made no reference to such habitat loss, although it is noted 
that none of this habitat is protected and that at one favoured site (30 km north of Toliara) 
some encroachment by villagers is occurring (Turner 1984). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED Full protection for all ground-rollers has been 
called for (Salvan 1970). Immediate protection of its habitat and protected area status for a 
representative tract of Didierea were formally recommended in 1970 (IUCN 1972). A study to 
determine the extent and type of habitat destruction reported in the Long-tailed 
Ground-roller's range (see under Threats) is urgently needed, together with a detailed 
ecological study of the bird to provide data essential to any strategy for its long-term 
conservation, particularly in relation to its apparent seasonal movements. Both this and the 
equally remarkable Subdesert Mesite, whose ranges are almost exactly coincident, merit 
conservation by means of a protected area. 

REMARKS This extraordinary bird occupies a monotypic genus. Concerning its occurence 
south of Toliara, a peak named "Mahinia ou Table" and some low hills named "Chaine de la 
Table" are indicated as lying behind the coast between the Fiherenana and Onilahy estuaries (in 
Locamus 1900). 



YELLOW-BELLIED SUNBIRD-ASITY INDETERMINATE 

Neodrepanis hypoxantha Salomonsen, 1933 Passeriformes: Philepittidae 

SUMMARY This Madagascar endemic species, difficult to distinguish from its only congener, 
is known from 12 specimens collected before 1930, although it was recognised as a species only 
in 1933. The generally held view that it is unlikely to be extinct, despite extensive forest 
destruction within its known range, is supported by the discovery of a nesting pair in 1976, but 
it must be rare at best. 

DISTRIBUTION The Yellow-bellied Sunbird-asity was recognised as a species distinct from 
the Wattled Sunbird-asity Neodrepanis coruscans only in 1933, and no specimens of it have 
been collected since; it is known only from eastern-central Madagascar, in forests east and 
perhaps south of Antananarivo, and the Sihanaka forest. Data from the 12 (or 13: see 
Remarks) currently known specimens, in order of their collection, are as follows: 



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Appendix 3: species accounts 

one male, no locality or date but prior to October 1879 (Benson 1971b); 

one male, central Madagascar, June 1880 (Eck 1968, Benson 1976a); 

three males, one female, Andrangoloaka, November 1880 (Stresemann 1937, 
Salomonsen 1965; also Eck 1968, Benson 1976a); 

two males, one female, east of Antananarivo, July 1881 (Salomonsen 1933, 
Benson 1974); 

one male, "E. Imerina" (near Antananarivo), October/November 1895 (Wetmore 
1953); 

one male, Sihanaka forest, 25 February, probably 1925 (Greenway 1967); 

one male, Fito (i.e. Sihanaka forest), August 1929 (Salomonsen 1965). 
Andrangoloaka was reported, at second-hand, to be situated high (1,000-1,300 m in one 
account, 1,400 m in another) on the eastern slopes of the plateau east of Antsirabe, c. 150 km 
south of Antananarivo, but this settlement and the great majority of surrounding forest no 
longer exist (Stresemann 1937, Greenway 1967). This information appears without doubt to be 
wrong, however, since there is or was an "Andrangolaoka" (sic) at 19°02'S 47°55'E on the upper 
slopes of the forested escarpment immediately east of the Mantasoa reservoir east of 
Antananarivo (Office of Geography 1955; not marked in IGNT 1964): in terms of the other 
records, this appears no less likely a locality than at Antsirabe, which is well outside the miin 
rainforest belt, and indeed it appears as both "Andrangaloaka" and "Andrangoloaka" on separate 
nineteenth century maps which show no similar name anywhere near Antsirabe (see Laillet and 
Superbie 1889, Locamus 1900; also Remarks). The forests around Antananarivo, from the 
eastern parts of which the type-material came (Salomonsen 1933), no longer exist (Salomonsen 
1934; also Wetmore 1953), nor do those at "East Imerina" (D. A. Turner pers. comm. 1983), but 
the species is expected to survive in Sihanaka forest, to the north-east of the Imerina plateau 
(Salomonsen 1965, Greenway 1967, Benson 1974). A stand of original forest at Tsinjoarivo, 
near Antsirabe, was twice identified as a likely site for the species (Lavauden 1937, Greenway 
1967; see Remarks), the remaining tiny patches of forest at the former locality of 
Andrangoloaka could perhaps still have held some birds (Stresemann 1937), and forest east of 
Anjozorobe (not Ankazobe as reported in King 1978-1979), north-east of Antananarivo, was 
also considered worth investigating (Lavauden 1937), but whether and in what condition these 
forests still survive is unknown. The Fierenana district north of Perinet has also been 
suggested as a possible site for the species (King 1978-1979). In December 1973 and 
November 1976 birds were seen and photographed (originals and copies with VIREO at ANSP) 
in forest several miles from and c. 100 m higher than the Perinet- Analamazaotra Special 
Reserve (A. D. Forbes-Watson pers. comm. 1984). 

POPULATION Numbers are unknown. While only nineteenth century records, all from the 
now largely deforested central parts of Madagascar, were known, the species was considered at 
best very rare and probably extinct, a judgement reinforced by the failure of the Mission 
Franco-Anglo-Americaine of 1929-1931 to find any (Salomonsen 1933,1934, Lavauden 1937, 
Stresemann 1937, Wetmore 1953). However, the discovery that birds had been collected in 
Sihanaka forest in the 1920s has resulted in confident predictions of its survival, albeit in low 
densities (Salomonsen 1965, Greenway 1967, Benson 1974), and these have been borne out by 
the 1976 sighting even though this was not itself in Sihanaka (see Distribution). The species 
was noted for being inexplicably uncommon in the last century around Andrangoloaka 
(Hildebrandt 1881; see Remarks). 

ECOLOGY This species inhabits rainforest; it is regarded as possibly a highland counterpart of 
the very similar Wattled Sunbird-asity (Wetmore 1953; also Lavauden 1937, A. D. 
Forbes- Watson pers. comm. 1984), although records from Sihanaka forest indicate possible 
sympatry between the two species (Benson 1974). It is a nectar-feeder like the Wattled 
Sunbird-asity; both species assume breeding plumage in September/November (Salomonsen 
1965). A pair observed near Perinet in December 1973 were feeding young in a (sunbird-like) 
nest placed 4-5 m up in thick forest (A. D. Forbes-Watson pers. comm. 1984). The species 
may inhabit canopy and thus have escaped notice (Benson 1980). However, observations a 
century ago refer to it feeding at flowering bushes in primary forest clearings, also to its call 
being a barely audible soft whistle (Hildebrandt 1881; see Remarks). In November 1976 near 
Perinet a male was observed feeding at Loranthus blossom; in December 1973 at the same 
locality a male was watched fly-catching alate termites and feeding them to nestlings (A. D. 
Forbes-Watson pers. comm. 1984). 

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An environmental profile of Madagascar 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. 

REMARKS Concerning the number of museum specimens of this species, there is apparently a 
thirteenth in Sydney, Australia, labelled merely "Antananarivo" (A. D. Forbes-Watson pers. 
comm. 1984). Along with two other eastern Madagascar forest birds, the Dusky 
Greenbul Phyllastrephus tenebrosus and the Red-tailed Newtonia Newtonia fanovanae, the 
Yellow-bellied Sunbird-asity is thought likely to have been overlooked owing to its sparseness, 
elusiveness and difficult habitat (Benson 1974). Its validity as a full species has recently been 
reaffirmed (Benson 1974). The Tsinjoarivo intended as a probable site for the species is most 
likely that at 19°37'S 47°40'E, i.e. on the edge of the main rainforest belt, and not that at 
19°54'S 46°39'E (in Office of Geography 1955). Observations in the last century (Hildebrandt 
1881), given under the name of the Wattled Sunbird-asity, appear to refer exclusively to the 
Yellow-bellied species, reference being made to "brilliant yellow" undersides, only a few 
specimens being collected (J. M. Hildebrandt was the collector of all four specimens from 
Andrangoloaka listed under Distribution), and (elsewhere in the paper) Andrangoloaka as the 
locality in which much fieldwork had been done (and where indeed the paper was written). 



APPERT'S GREENBUL RARE 

Phyllastrephus apperti Colston, 1972 Passeriformes: Pycnonotidae 

SUMMARY This ground-haunting, dry forest bulbul is known with certainty from only two 
remote unprotected localities in south-west Madagascar, where it is exceptionally rare and 
faces the danger of destruction of its habitat by fire. 

DISTRIBUTION Appert's Greenbul was first found in a forest 40 km south-east of 
Ankazoabo, south-west Madagascar, where it was twice seen, on 7 June and 4 September 1962; 
on the latter date two specimens were collected, from which the species was described ten years 
later (Colston 1972). Throughout this intervening period the collector never found the bird 
again during studies over an area of 40,000 km 2 around the Mangoky River (Colston 1972). 
On 20 August 1974 it was rediscovered in the same patch of forest as in 1962 and a specimen 
was collected and sent to Antananarivo (O. Appert in lilt. 1983; for numbers then seen, see 
Population). A single bird was seen in the nearby Zombitsy forest, east of Sakaraha, in July 
1974 (Benson and Irwin 1975), two or three were present the following month (O. Appert in 
lilt. 1983), and birds have been consistently seen there subsequently (see below). Two 
specimens, probably of this species, were collected east of Toliara ("probably near Sakaraha") 
and were deposited for a time in Antananarivo, but are now lost (Colston 1972; but see under 
Distribution and Remarks for Grey-crowned Greenbul Phyllastrephus cinereiceps). The sites 
south-east of Ankazoabo and east of Sakaraha belong to (what was at least in 1963) a fairly 
unfragmented (but variously named) block of forest (Vohibasia forest, Jarindrano forest, 
Mangona forest, Zombitsy forest), extending to the north and south of Andranolava and very 
roughly covering 1,000 km 2 (see IGNT 1964). Nevertheless, despite fieldwork throughout the 
Zombitsy forest, 1976-1981, the species was always only ever found in the same single area in 
one corner, only 0.5 km 2 in size (D. A. Turner pers. comm. 1983). A recent (but undated) 



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Appendix 3: species accounts 

record of one (seen on the ground) "15 km east of Sakaraha" (O. Langrand in litt. 1984) perhaps 
involves a new (third) site for the species. 

POPULATION Numbers are probably very small. Soon after its first description it was 
considered possibly quite common (Forbes- Watson el al. 1973), but subsequent evidence is 
against this: apart from what is said above under Distribution, it could not be relocated by one 
observer in the Zombitsy forest in August 1974 (Benson and Irwin 1975) nor was it found at 
another nearby site east of Sakaraha during five days' fieldwork, 1972-1973 (Benson 1976b). 
From this it would appear that the species is very local and sparse within its small known 
range. At the corner of the Zombitsy forest where it could always be found, 1976-1981, seven 
to eight birds were usually to be seen, though once 15; probably 20-30 birds is the maximum 
for this area (D. A. Turner pers. comm. 1983). In the forest south-east of Ankazoabo on the 
last occasion the species was seen there (20 August 1974), two groups were encountered, one o f 
two to three and the other of about eight birds (O. Appert in litt. 1983). 

ECOLOGY The species inhabits dense dry forest, searching for food in the leaf-litter (Colston 
1972). Although it is reported to be highly terrestrial, behaving rather like an 
akalat Sheppardia, rarely moving more than 1 m above ground, and always occurring in groups 
of five to eight (Colston 1972, D. A. Turner pers. comm. 1983), the bird seen in Zombitsy 
forest in July 1974 was single (though in the company of c. 20 Long-billed 
Greenbuls Phyllastrephus madagascariensis) and kept 1-5 m above ground (Benson and Irwin 
1975): indeed, observations in Zombitsy forest in November 1976 were commonly of birds 
clinging to liana tangles up to 2 m from the ground, in the manner of reed 
warblers Acrocephalus (A. D. Forbes-Watson pers. comm. 1984). Birds have once been seen to 
go higher in the trees when disturbed, and they may do so habitually at times when the forest 
is more humid (O. Appert in litt. 1983). The Zombitsy forest is much greener in the dry season 
than that south-east of Ankazoabo, retaining some of its leaves (O. Appert in litt. 1983). 

THREATS The highly restricted range of this species must be a source of permanent concern 
and vigilance for its welfare, and in late 1978 or early 1979 a cyclone destroyed forest as close 
as 2 km to the single known area in Zombitsy (D. A. Turner pers. comm. 1983); moreover, 
forest burning in this region has apparently been very serious in recent years (O. Appert in litt. 
1983). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED Further fieldwork to determine the range and 
status of this species is desirable, but protection of the forests from which it is known is 
perhaps more immediately important. Control of the present forest cutting and burning is 
essential for ecological stability in the region (O. Appert in litt. 1983). 

REMARKS Study of the Antananarivo specimen, alongside the type, is desirable for absolute 
confirmation of the validity of the species (Colston 1972). 



DUSKY GREENBUL RARE 

Phyllastrephus tenebrosus (Stresemann, 1925) Passeriformes: Pycnonotidae 

SUMMARY This mysterious bulbul of rainforest undergrowth is known from only eight skins 
and two adjacent localities (Sihanaka forest and Perinet-Analamazoatra), eastern central 
Madagascar. 

DISTRIBUTION The Dusky Greenbul was first described from four specimens collected in 
the Sihanaka forest, eastern Madagascar, of which one was collected in December 1924 (type), 
one in June 1925 and one in December 1925 (Stresemann 1925, Benson el al. 1976, 
D. K. Read in lilt. 1983). Two further specimens from Sihanaka forest (undated) were 
acquired by the Mission Franco-Anglo-Americaine from Herschell-Chauvin around 1930 

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An environmental profile of Madagascar 

(Delacour 1932a, Rand 1936), and another was obtained there on 7 April 1929 (Lavauden 
1932). A bird was also shot north of Analamazaotra (near Perinet) on 8 May 1929 (Lavauden 
1932). A single bird was seen at Perinet on 25 June 1974, one on 23 November 1976 and one 
on 14 November 1977 (D. A. Turner in litt. 1983; also Benson and Irwin 1975). That the 
species occurs thoughout "forests in the east of Madagascar" (Lavauden 1937) appears an 
unacceptably sweeping assumption; that it may occur at other sites in the central section of 
these forests (Milon el al. 1973) seems a reasonable hope. 

POPULATION Numbers are unknown. This species probably goes unrecorded by combining 
extreme elusiveness and difficult habitat with general sparseness (Benson 1974). Its rarity has 
been remarked (Delacour 1932b, Lavauden 1932). 

ECOLOGY From existing records (see Distribution) it is confined to humid rainforest. It was 
judged probably a bird of ground-cover (Rand 1936) and the first observation in life was of a 
single bird with a pair of White-throated Oxylabes Oxylabes madagascariensis, moving 
through branches c. 2 m above the ground, presumably gleaning insects though it was not 
actually seen to feed (Benson and Irwin 1975). Subsequent observations confirm it to be an 
undergrowth species (D. A. Turner pers. comm. 1983). The bird seen in November 1976 was 
clinging to vertical stems 1-2 m from the ground (A. D. Forbes-Watson pers. comm. 1984). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). 

CONSERVATION MEASURES TAKEN The species occurs in the Perinet-Analamazaotra 
Special Reserve, which covers 810 ha (Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. 

REMARKS The Dusky Greenbul has been treated as a race of the Madagascar 
Greenbul Phyllastrephus madagascariensis (Milon et al. 1973) but its validity as a full species 
has since been reaffirmed (Benson 1974, Benson et al. 1976; also Benson and Irwin 1975). 



GREY-CROWNED GREENBUL RARE 

Phyllastrephus cinereiceps (Sharpe, 1881) Passeriformes: Pycnonotidae 

SUMMARY This Madagascar bulbul, probably confined to rainforest, remains almost totally 
unknown, and apparently has been found only twice in the past 50 years. 

DISTRIBUTION The Grey-crowned Greenbul possibly occurs throughout the rainforests of 
eastern Madagascar, but is known from only a few scattered sites. It was first described from 
Fianarantsoa, in the southern half of the eastern rainforest belt (Sharpe 1881), and subsequently 
found in the nearby Ankafana (= Tsarafidy) forest (seven specimens in BMNH, all March 1881: 
NJC; see Remarks), Sihanaka forest and at Fanovana (Delacour 1932a, Rand 1936), though this 
last area is now cleared (D. A. Turner pers. comm. 1983), and in the Tsaratanana massif (Milon 
1957). An observation of the species is reported from "Lambomakandro forest" to the east of 
Sakaraha in south-west Madagascar (Milon et al. 1973), but for a bird previously known only 
from the humid forest area of the island this record appears somewhat anomalous (see 
Remarks); if this is discounted, the species has only been recorded twice in the past 50 years, 



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Appendix 3: species accounts 

in the Tsaratanana massif (see above) and near Didy, on the western edge of the Sihanaka 
forest, in May 1971 (A. D. Forbes-Watson pers. comm. 1984). 

POPULATION The species was considered uncommon in Sihanaka forest 50 years ago 
(Delacour 1932a) and is known from apparently only a single specimen from Tsaratanana 
(Milon 1957). 

ECOLOGY It inhabits the ground-cover of deep rainforest, gleaning for insects through the 
low bushes, and associating with the Short-billed Greenbul Phyllastrephus zosterops (Rand 
1936). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). 

CONSERVATION MEASURES TAKEN The species presumably occurs in the Tsaratanana 
Nature Reserve (R.N.I, no. 4), which covers 48,622 ha (Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. The forest at Tsarafidy where this species occurs 
was identified in 1961 as a place of exceptional interest for its wildlife which certainly 
deserved complete protection (Griveaud 1961); the species may be expected in the nearby 
"Nandehizana" forest, if this still survives (for both localities see Conservation Measures 
Proposed and Remarks under Madagascar Yellowbrow Crossleyia xanthophrys). 

REMARKS It is possible that "Fianarantsoa" was a generalised locality and that the 
type-material actually came from (the fairly nearby) Tsarafidy forest. The anomalous record 
of this species from east of Sakaraha, south-west Madagascar, given (in Milon et al. 1973) 
without any further information, may possibly be connected with the two lost specimens 
thought to be of Appert's Greenbul Phyllastrephus apperti, which came from "east of Tulear, 
probably near Sakaraha" (Colston 1972; see relevant sheet): two villages named Lambomakandro 
are situated in the northern parts of the Zombitsy forest (in IGNT 1964), which is one of only 
two only certain localities for Appert's Greenbul. 



VAN DAM'S VANGA RARE 

Xenopirostris damii Schlegel, 1866 Passeriformes: Vangidae 

SUMMARY This insectivorous bird of deciduous forest is known this century from a single 
site (Ankarafantsika) in north-west Madagascar which is, however, a protected area. 

DISTRIBUTION Van Dam's Vanga was originally described from two specimens collected on 
9 October 1864 in the forests near Ambassuana (Ambasohana) in the far north-west of 
Madagascar (Schlegel 1866, Schlegel and Pollen 1868). At least six further specimens were 
collected at around this time and probably at this locality (given as Pasandava Bay) (Milne 
Edwards and Grandidier 1885). The generalisation of the type-locality as Pasandava 
(Ampasindava) Bay (south of Nosy Be) (e.g. Milne Edwards and Grandidier 1885, Lavauden 
1932,1937, Milon el al. 1973) or else as "the Sambirano" (Benson et al. 1977), i.e. the general 
region of the river of that name which runs into this bay, is misleading: according to the 
contemporary map (in Pollen 1868) Ambassuana lay on the river of the same name, some 
20 km east of the easternmost part of Pasandava Bay and much closer to Ambara (Ambaro) 
Bay: it would appear to have been situated at about 13°35'S 48°40'E and a few kilometres east 

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An environmental profile of Madagascar 

or south of the locality now called Maherivaratra (past which the Ambazoana River flows), and 
indeed there is an "Ambazoana Bala" marked at roughly this spot in a 1900 map (see Locamus 
1900). There have been no further records of the species from the far north-west, but it 
appears that the Ambazoana valley has never been revisited. The species was rediscovered on 
9 October 1928 when a male was collected on the Ankarafantsika plateau, south-east of 
Mahajanga, and a juvenile was collected there on 5 July 1930 (Lavauden 1932); the species was 
found and collected there again in 1969-1971 (Salvan 1970, Forbes-Watson et al. 1973) and has 
been regularly seen subsequently (D. A. Turner pers. comm. 1983, O. Langrand in litt. 1984); 
the area involved in the post- 1969 observations is near Lake Ampijoroa (Milon et al. 1973, 
A. D. Forbes-Watson pers. comm. 1984; see Remarks). 

POPULATION Although this species was unanimously described as "very rare" (Schlegel and 
Pollen 1868, Delacour 1932a, Lavauden 1932) and even treated as Endangered in King 
(1978-1979), the most recent observations at Ankarafantsika indicate that it is in fact present 
in fairly good numbers there, with certainly 50 or more pairs in one relatively small area where 
the forest is untouched (D. A. Turner pers. comm. 1983). 

ECOLOGY It inhabits primary deciduous forest (Delacour 1932a, D. A. Turner pers. comm. 
1983); it is seen along the edges of woods foraging for insects (Schlegel and Pollen 1868). The 
stomachs of the first two known specimens contained beetle remains (Schlegel and Pollen 
1868). Although described as solitary (Schlegel and Pollen 1868) it was also reported to occur 
in small groups of six to eight birds (Milne Edwards and Grandidier 1885). 

THREATS The highly restricted range of this species must be a source of permanent concern 
and vigilance for its welfare. Deforestation is likely to have affected many areas where it 
might have been searched for in north-west Madagascar. 

CONSERVATION MEASURES TAKEN The area where it has been recorded this century 
partly falls within the Ankarafantsika Nature Reserve (R.N.I, no. 7) (see Andriamampianina 
1981). 

CONSERVATION MEASURES PROPOSED A study of the status and ecology of this bird at 
Ankarafantsika would help determine where else it might be searched for and what 
management it might require (besides confirming, for example, that it is resident throughout 
the year). An investigation of the type-locality (see Distribution) is warranted to establish 
whether any original tracts of forest remain and whether they still hold populations of this 
species. All such work should be undertaken in conjunction with studies recommended under 
Conservation Measures Proposed for the White-breasted Mesite Mesitornis variegata. 

REMARKS Although this species has been considered doubtfully distinct from Lafresnaye's 
Vanga Xenopirostris xenopirostris (see, e.g., Delacour 1932a, Appert 1970) and confused with it 
(see, e.g., Bartlett 1875, Menegaux 1907), it has been affirmed as a good species (in Lavauden 
1932) and this judgement is accepted here. The importance of the Ankarafantsika Nature 
Reserve as the only locality currently known for the species and the White-breasted Mesite 
cannot be overstated. However, it is to be noted that (according to IGNT 1964) Ampijoroa is 
well outside the boundaries of the nature reserve, and right next to the main road from 
Antananarivo to Mahajanga. 



POLLEN'S VANGA RARE 

Xenopirostris polleni (Schlegel, 1868) Passeriformes: Vangidae 

SUMMARY This insectivorous rainforest bird is known from a wide variety of localities in 
eastern Madagascar but is everywhere rare. 

DISTRIBUTION The scatter of records for Pollen's Vanga indicates that it is confined to the 
rainforests of eastern Madagascar. However, the type-locality is the "north-west coast" 

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(Schlegel and Pollen 1868) (attempts to trace anything more precise have been fruitless). If the 
bird is confined to humid forest, and if the map of the area explored by the original collectors 
(in Pollen 1868) represents the total area they prospected, then (according to the vegetation 
map in Rand 1936) the only suitable localities in the north-west are the Montagne d'Ambre or 
the western parts of the Tsaratanana massif, particularly the Manongarivo massif (west of the 
Sambirano) (see IGNT 1964). The original three specimens were later described as coming 
from "north-east" Madagascar (Hartlaub 1877) but this is evidently in error (see Remarks). 
However, there is a suspected sight record from Marojejy (north-east) in September 1972 
(Benson et al. 1977) and one was seen at 500 m near Maroantsetra in November 1982 
(O. Langrand in litt. 1984). All other records except one are from the central parts of eastern 
Madagascar, listed here from north to south. There are two specimens in BMNH from 
Fenoarivo Atsinanana, dated May 1895 (Benson et al. 1977, NJC), and twelve in MNHN and 
MRAC from Sihanaka forest (NJC; also Lavauden 1932), one of these latter actually being 
taken between Fanovana and Beforona (see Remarks). Specimens are known from both 
Fanovana (c. 800 m), April 1931 (Delacour 1932a, Rand 1932,1936) and August 1932, and 
Beforona, September 1932 (specimens in Stockholm: C. Edelstam in litt. 1983), but forest at 
least at the former locality no longer exists (D. A. Turner pers. comm. 1983). Individuals have 
been seen in November 1976, July 1980 and December 1982 at Perinet (D. A. Turner in litt. 
1983, O. Langrand in litt. 1984). Several specimens were reported to have been collected in the 
forests on the eastern slopes of the great central massif (Milne Edwards and Grandidier 1885), 
although the only traceable reference to any of these is to one from south-east of 
Antananarivo, February 1872 (Sharpe 1872; see Remarks); however in UMZC there are three 
specimens labelled "Imerina, 1891" (Imerina is the whole area around Antananarivo: see 
Deschamps 1960) and another taken within 60 km of Antananarivo, 1881 (NJC), and 
presumably all four were from "the forests of the eastern slopes" in central Madagascar. Single 
adult males were collected at Mahanoro on 1 May and the "Sakales" River (see Remarks) on 
15 May 1895 (Richmond 1897). In 1959 a specimen was collected in Tsarafidy forest between 
Ambohimahasoa and Fianarantsoa (Griveaud 1961), two specimens were collected there ("Foret 
d'Amboasary") in January 1961 (Benson et al. 1977), two birds were seen (one mist-netted) 
there in April 1971 (Forbes-Watson et al. 1973, D. A. Turner in litt. 1983), and a specimen in 
SMF labelled as Lafresnaye's Vanga Xenopirostris xenopirostris and taken at "Amboasary" on 
27 October 1931 (NJC) presumably also originates from Tsarafidy. In MNHN there is also a 
specimen from 30 km north-north-west of Taolanaro in the far south-east of Madagascar, 
collected on the 26 May 1948 (NJC). 

POPULATION Although unanimously regarded as rare (Delacour 1932a, Rand 1936, Milon et 
al. 1973) and even as Endangered (in King 1978-1979), it is clear from the evidence above that 
this species has a much wider range than has often been stated. However, the nineteenth 
century report of its occurrence in groups (see Ecology) has not been confirmed, which might 
(conceivably) indicate a greater decline in numbers than that resulting simply from the decline 
in total forest area. It is evidently only ever present in small numbers wherever it survives. 

ECOLOGY Pollen's Vanga is apparently a bird of primary rainforest. It was reported 
(presumably by a collector in a personal communication) to live in groups of 8-10, to feed on 
insects, small reptiles and frogs, to fly low and to be tame (Milne Edwards and Grandidier 
1885). The MNHN specimen from Taolanaro contained a very large spider, a caterpillar and 
insect remains; it was not in breeding condition (NJC). All sight records given under 
Distribution (from Maroantsetra, Perinet and Tsarafidy as well as the possible one from 
Marojejy) were of birds in mixed bird-parties (Benson et al. 1977, D. A. Turner in litt. 1983, 
O. Langrand in litt. 1984), so it is possible that the groups of 8-10 referred to in the last 
century were not intended to imply monospecific flocks. 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). It is not known whether primary forest still occurs in the 
coastal areas where the species was collected in the last century (Fenoarivo Atsinanana, 
Mahanoro, "Sakales" River), but this seems unlikely. 

CONSERVATION MEASURES TAKEN The species occurs in the Perinet-Analamazaotra 
Special Reserve, which covers 810 ha, and possibly in the Marojejy Nature Reserve (R.N.I. 

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no. 12), which covers 60,150 ha (Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sihanaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. The forest at Tsarafidy where this species occurs 
was identified in 1961 as a place of exceptional interest for its wildlife which certainly 
deserved complete protection (Griveaud 1961); the species may be expected in the nearby 
"Nandehizana" forest, if this still survives (for both localities see Conservation Measures 
Proposed and Remarks under Madagascar Yellowbrow Crossleyia xanthophrys). 

REMARKS The error concerning the type-material originating from north-east Madagascar 
arises from the fact that the specimen-labels say "N. O. Madagascar": in both Dutch and 
German this would signify north-east, and was taken as such by Hartlaub (1877), but the 
language used on the labels is French ("voyage de Van Dam"), hence "N. O." signifies 
north-west (specimens in RMNH: NJC). That one of the specimens in MNHN labelled as from 
Sihanaka should also say "route de Fanovana a Beforona" (i.e. part of the road between 
Antananarivo and Toamasina) greatly stretches the limits accepted here of the "Sihanaka forest" 
(see Conservation Measures Proposed under Madagascar Serpent Eagle). "Kinkimauro" was 
given as the precise locality of the specimen from south-east of Antananarivo (Sharpe 1872), 
and later repeated as "Kinkimanro" (Hartlaub 1877), but "kinkimavo" was in fact a widespread 
native name for certain grey birds (Milne Edwards and Grandidier 1885) and it is obvious that 
the word was written on the specimen's label by the collector and misinterpreted as a site by its 
recipient. The "Sakales" river cannot be traced but it is evident that the collector was very 
close to the Sakaleona River (south of Mahanoro) at the time and these are doubtless identical 
("Sakales" is obviously a misreading of Sakaleo, a village which in the last century stood at the 
estuary of the river: see Locamus 1900); both this and the site at Mahanoro have been included 
in a map of the species's distribution (see Benson et al. 1977) although there is no reference to 
Richmond (1897) as the source. There is a striking similarity between the plumages of the 
Tylas Vanga Tylas eduardi and immature Pollen's Vanga (Sharpe 1870, Benson 1971b, 
Benson et al. 1977, O. Langrand in litt. 1984) and, as it appears that the two species are 
sympatric and possibly without differences in habitat, a field study of their relationship has 
been urged (Benson et al. 1977). 



BENSON'S ROCKTHRUSH INSUFFICIENTLY KNOWN 

Monticola bensoni Farkas, 1971 Passeriformes: Muscicapidae 

SUMMARY This recently described rock-haunting thrush is possibly quite widespread but as 
yet is known only from a few dry rocky areas in south-west Madagascar. 

DISTRIBUTION Benson's Rockthrush was recently described from two old specimens 
collected by "Zaast" at an unknown time in an intraceable locality ("Ankarefu, Antinosy Cy") in 
"south-west" Madagascar (Farkas 1971). In 1962 birds were discovered and recognised as a new 
species at several localities in the Mangoky River region and the northern Isalo massif, and 
found again in 1969 and 1970 in the latter locality and at a single site between Ihosy and 
Zazafotsy (east of the northern Isalo massif) (Farkas 1971). In August 1969 and June 1971 the 
species was found 150 km south of the northern Isalo massif (D. A. Turner in litt. 1983). On 
10 July 1977 two birds were observed on telephone wires along the main road running through 
the Zombitsy Forest Reserve near Sakaraha (D. A. Turner in litt. 1983). Excluding the records 
from Antinosy County (southern Madagascar; but see Remarks), all sightings are from between 
the Mangoky and Onilahy Rivers, south-west Madagascar, but the distribution of the species 
has been anticipated as covering one-fifth of the island, from "at least" the eastern fringes of 
the Bemaraha plateau in the central-west to "Antinosy County" in the south (see map in Farkas 

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Appendix 3: species accounts 

1971, also under Remarks); it should however be noted that localised distribution, despite more 
extensive and apparently similar habitat, is a characteristic of several threatened Madagascar 
birds (e.g. Subdesert Mesite Monias benschi, Appert's Greenbul Phyllastrephus apperti) and a 
wider range for this species cannot be assumed with confidence. In the Mangoky region birds 
were only seen in winter (June/July) and in different habitat (see Ecology), and it seems likely 
that some local migration occurs at this season (Farkas 1971). 

POPULATION Numbers are unknown. At least six males (some paired) were found along a 
2 km stretch of road in one locality (Farkas 1971), so the population density is probably fairly 
high and stable in suitable habitat. Nevertheless, on present information it remains possible 
that total numbers are very small (Forbes-Watson et al. 1973). 

ECOLOGY This is the only rock-inhabiting Monticola in Madagascar, apparently preferring 
huge rocks and extended cliff faces rising steeply out of open rolling hillsides, birds keeping 
mostly to rocks, occasionally visiting thin bushes, small trees, etc., but retreating to high rocks 
in alarm; food is insects sometimes caught in flight (Farkas 1971). Display-flights are 
performed from the highest available rock-peaks, the song being loud, clear, attractive; 
territories may be as close as 200 m and the species is probably double-brooded or else a late 
summer breeder (Farkas 1971). In winter, some birds are found in dry riverbeds, rocky 
canyons, hill slopes with huge boulders, all with scattered bushes and trees (Farkas 1971); the 
birds on wires by the roadside in dry forest at Zombitsy were also presumably wintering 
(D. A. Turner in lilt. 1983). 

THREATS None is known, but the population could prove to be extremely low and 
restricted. 

CONSERVATION MEASURES TAKEN The Isalo massif is protected as a national park 
(Andriamampianina 1981). It is to be noted that the original authority did not collect any new 
specimens but used one of the two old skins for the type (Farkas 1971). 

CONSERVATION MEASURES PROPOSED Studies are needed to determine the extent of this 
species's breeding and wintering range, undertaken in the course of other fieldwork in the 
general region of and at increasing distances from the Isalo massif. 

REMARKS Although treated as a race of the "Madagascar Rockthrush" Pseudocossyphus 
imerinus (Milon et al. 1973), the status of Benson's Rockthrush as a valid species has been 
reaffirmed (see Benson et al. 1977). The map in Farkas (1971) shows "Antinosy [sic] County" 
in the far south of Madagascar; however, maps in Deschamps (1960) show that the Antanosy 
people occupied and still occupy the extreme south-east part of southern Madagascar, south 
and east of the area shown by Farkas. While it is still possible that Benson's Rockthrush occurs 
in the southern area shaded by Farkas (1971), e.g. in the Ivakoany massif, the maps in 
Deschamps (1960) show an area of south-west Madagascar, across the Onilahy River at the 
southernmost end of the Isalo massif, colonised by "Antanosy emigres", and this seems more 
likely to be the type-locality of the species (the "Cy" on the type's specimen-label probably 
specifies "country" rather than the assumed "County", hence something less institutionalised and 
fixed). Moreover, of 1 1 gazetteered localities under the name "Ankarefo" ("Ankarefu" is not 
listed) in Office of Geography (1955), only one falls into either area of Antanosy people as 
marked by Deschamps, this being just north of Betroka at 23°06'S 46°06'E, some 100 km east 
of the Isalo massif. 



MADAGASCAR YELLOWBROW INDETERMINATE 

Crossleyia xanthophrys (Sharpe, 1875) Passeriformes: Muscicapidae 

SUMMARY This distinctive species is confined to Madagascar's rainforests, where it has been 
seen only twice in the past 50 years. 



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DISTRIBUTION The Madagascar Yellowbrow is known from rainforest in central eastern 
Madagascar, with one record from the north. It was first described from a single bird collected 
on the "east coast" of Madagascar (Milne Edwards and Grandidier 1881). Subsequently it was 
widely collected but very poorly documented in the literature. Published records before 1930 
are of three specimens from Fianarantsoa received in BMNH in 1880.(Sharpe 1881; NJC), the 
species as common in Ankafana (= Tsarafidy) and "Nandehizana" forests, also around 1880 
(Deans Cowan 1882; see Remarks), a specimen presumably from the eastern Imerina forests in 
the 1890s (Oberholser 1900), and a "considerable number of skins" (at least eighteen) procured 
by one collector from Sihanaka forest (Delacour 1932b, Rand 1932). In BMNH there are in 
fact six specimens from Ankafana (which may well be the locality intended by "Fianarantsoa", 
which is fairly nearby) and six also from Sihanaka, in MNHN there are 12 from Sihanaka, and 
in BMNH and MRAC together there are also two from Brickaville district, 1925, six from 
Andevoranto forest, 1925, four from Lakato forest, 1924-1925, and one from "Betsileo", 
undated (NJC,SNS), "Betsileo" indicating south-central Madagascar (see map in Deschamps 
1960, also Locamus 1900). Despite this relative wealth of records, since 1930 there have been 
only two: one in the Tsaratanana massif in the north in 1966 (Albignac 1970) and one at Perinet 
east of Antananarivo in July 1968 (Benson and Irwin 1975). 

POPULATION Along with the Yellow-bellied Sunbird-asity Neodrepanis hypoxantha, Dusky 
Greenbul Phyllastrephus tenebrosus and Red-tailed Newtonia Newtonia fanovanae, this species 
is considered difficult to observe because of its elusiveness, sparseness, and difficult habitat 
(Benson 1974, Benson and Irwin 1975). That one particular collector should have obtained a 
"considerable number" of specimens probably only reflects the time-period over which his 
collections were made (40 years in one account: see Distribution under Madagascar Red 
Owl Tyto soumagnei) and, perhaps, the hunting techniques of the natives from whom most of 
his ornithological material was acquired (see Rand 1932). 

ECOLOGY General "habits" were considered likely to prove similar to the Grey-crowned 
Oxylabes Oxylabes cinereiceps, which is a bird of ground-cover in heavy forest and gleans for 
insects through low bushes (Rand 1932). The bird seen in 1968 was in undergrowth on the 
edge of evergreen forest (Benson and Irwin 1975). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). The forest in eastern Imerina is now all cleared (D. A. Turner 
pers. comm. 1983), and it seems unlikely that forest near the coast at Brickaville or 
Andevoranto would now be extant. 

CONSERVATION MEASURES TAKEN The species has been recorded from the Tsaratanana 
Nature Reserve (R.N.I, no. 4), which covers 48,622 ha, and the Perinet-Analamazaotra Special 
Reserve, which covers 810 ha (Andriamampianina 1981). 

CONSERVATION MEASURES PROPOSED Immediate and effective protection of as much 
remaining rainforest as possible would almost certainly guarantee the survival of this and all 
other rainforest-dependent species in Madagascar; and at least, on current knowledge, complete 
protection of the intact parts of Sianaka forest is of extreme importance (see Conservation 
Measures Proposed under Madagascar Serpent Eagle). Any ornithological work in the other 
areas from which the species is known, or where it might be expected, should where possible 
be extended to include searches to locate it. The forest at Tsarafidy where this species occurs 
was identified in 1961 as a place of exceptional interest for its wildlife which certainly 
deserved complete protection (Griveaud 1961); protection for "Nandehizana" forest, if it 
survives, is also merited, and a survey to relocate and evaluate this forest is warranted (see 
Remarks). 

REMARKS Tsarafidy (i.e. Ankafana) forest, north of Fianarantsoa, is important not only for 
this species but also for Pollen's Vanga Xenopirostris polleni and the Grey-crowned 
Greenbul Phyllastrephus cinereiceps (see relevant accounts), Pitta-like Ground-roller Atelornis 
pittoides and Brown Emu-tail Dromaeocercus brunneus, as well as many rare lemurs, 
invertebrates and plants (Deans Cowan 1882, Griveaud 1961). The location of "Nandehizana" 
forest, also of substantial importance (see Deans Cowan 1882), has not hitherto been traced, but 

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Appendix 3: species accounts 

there is a Nandihizina marked on an old map (see Locamus 1900) somewhat to the north of 
what is now called Tsarafidy, and which on a modern map would be located at 20°47'S 
47°10'E, i.e. the block of forest straddling the road between Ambositra and Ambohimahasoa 
(in IGNT 1964). A taxonomic reassessment of this bird has judged it to be a babbler 
(Timaliinae) not a bulbul (Pycnonotidae) (Benson and Irwin 1975), and it now reoccupies the 
monotypic genus created for it in 1877 (Hartlaub 1877), being aberrant enough possibly to 
require placing in a separate tribe, the Crossleyini (Irwin 1983). 



RED-TAILED NEWTONIA INDETERMINATE 

Newtonia fanovanae Gyldenstolpe, 1933 Passeriformes: Muscicapidae 

SUMMARY This flycatcher is known only from a single specimen from a forest, now cleared, 
in eastern central Madagascar. If it is not an invalid taxon based on an aberrant bird, it is 
either greatly overlooked, genuinely rare, or extinct. 

DISTRIBUTION The type and only specimen (adult, sex unknown) of the Red-tailed 
Newtonia was collected in the Fanovana forest, eastern central Madagascar, in December 1931 
(Gyldenstolpe 1933). Although its late discovery may reflect a very limited area of distribution 
(Gyldenstolpe 1933) it probably inhabits "the forest of the central part of the Humid East" 
(Rand 1936) and may occur in the Sihanaka forest (Salomomsen 1965) and at Perinet (Benson et 
al. 1977), although observations at Perinet in recent years have failed to find it (D. A. Turner 
pers. comm. 1983). 

POPULATION Numbers are unknown, but the species is regarded as probably sparse (Benson 
1974). 

ECOLOGY The ecology of this species is wholly unknown, other than that it must, as a 
flycatcher, be insectivorous. It has been speculated, on the basis of the plumage and ecological 
characters of its congeners, that the species may frequent the canopy of evergreen forest, in 
which it could be easily overlooked, especially if in any case uncommon (Benson 1974, 
Benson et al. 1977). Such speculation matches the contention that the species is a mimic of the 
female Red-tailed Vanga Calicalicus madagascariensis (see Remarks), since the latter is a bird 
of forest canopy (Milon et al. 1973, Benson et al. 1977). 

THREATS Destruction and disturbance of primary rainforest is the single most serious threat 
to this and all other rainforest-dependent species in Madagascar (see Threats under Madagascar 
Serpent Eagle Eutriorchis astur). The forest at Fanovana is now completely cleared (D. A. 
Turner pers. comm. 1983). 

CONSERVATION MEASURES TAKEN None is known. 

CONSERVATION MEASURES PROPOSED Protection of as much remaining rainforest as 
possible is the primary need; further study at certain sites, especially Perinet, to establish the 
continued existence and likely requirements of this species, is desirable. 

REMARKS Not having been seen for over 50 years, by CITES criteria the Red-tailed 
Newtonia could be treated as extinct. However, along with two other eastern Madagascar 
forest birds, the Yellow-bellied Sunbird-asity Neodrepanis hypoxantha and the Dusky 
Greenbul Phyllastrephus tenebrosus, it is thought likely to have been overlooked owing to its 
sparseness, elusiveness and difficult habitat (Benson 1974, Benson et al. 1977). Its validity as a 
full species has recently been reaffirmed (Benson et al. 1977), the possibility that it is an 
aberrant female Red-tailed Vanga (Forbes-Watson et al. 1973) being rejected on the grounds of 
its slender Newtonia bill, the Vanga having a short, stout bill and also differing in its larger, 
heavier size, black lower mandible, pale lores, and conspicuous broad (not narrow) white 
eye-ring (G. S. Keith in litt. 1983); nevertheless, the similarity in the plumage between the two 
is so "incredibly close. ..that this must be a case of mimicry" (C. Edelstam in litt. 1983). 

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Brit. Orn. Club 101: 240-241. 
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553-557. 
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van Someren, V. D. (1947) Field notes on some Madagascar birds. Ibis 89: 235-267. 



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Appendix 3: species accounts 



APPENDIX 3.B. MAMMALS 

Accounts for all lemur species and for the subspecies Lemur macaco macacao and L. macaco 
flavifrons are included. Tentative IUCN categories for each taxon are listed below; some of 
these will certainly require revision as more detailed information on the status of lemurs 
becomes available. 



E Allocebus trichotis 

nt Cheirogaleus major 

nt Cheirogaleus medius 

K Microcebus coquereli 

nt Microcebus murinus 

nt Microcebus rufus 

K Phaner furcifer 

K Avahi laniger 

E Indri indri 

K Propithecus verreauxi 

V Propithecus diadema 

E Daubentonia madagascariensis 

K Hapalemur griseus 

E Hapalemur simus 

K Lemur catta 



K Lemur coronatus 

nt Lemur fulvus 

V Lemur macaco macaco 

E Lemur macaco flavifrons 

V Lemur mongoz 

I Lemur rubriventer 

K Lepilemur dorsalis 

K Lepilemur edwardsi 

K Lepilemur leucopus 

K Lepilemur microdon 

K Lepilemur mustelinus 

K Lepilemur ruficaudatus 

K Lepilemur septentrionalis 

I Varecia variegata 



HAIRY-EARED DWARF LEMUR 



Allocebus trichotis (Giinther, 1875) 



Primates: Cheirogaleidae 



SUMMARY Endemic to Madagascar. Considered the rarest of all the lemurs and known from 
only four specimens collected from the eastern forests; three were collected last century and the 
fourth in 1965. Cause of decline unknown. Protected by law but not known to occur in any 
reserve. Any conservation measures will have to await the discovery of living specimens. 

DISTRIBUTION Madagascar. Known only from four specimens. The only 'modern' 
specimen was caught by A. Peyrieras in 1965 in the Andranomahitsy Forest to the west of 
Mananara on the east coast (4,9). The four records do suggest, however, that at one time the 
species occurred quite widely in the eastern humid forests (9). For map see (9). 

POPULATION It is possible that this animal is extinct; if not, the number remaining might be 
very low (4,8). However, Petter has recently commented that the species may be a cryptic 
canopy dweller and thus could be overlooked (10). An effort to locate the species in the 
Andranomahitsy Forest in 1975 was unsuccessful (6). The species is considered the rarest of 
the lemurs (9). 

HABITAT AND ECOLOGY Nothing known. The Andranomahitsy Forest is typical high rain 
forest of eastern Madagascar. 

THREATS Unknown. In 1972 Petter reported that habitat alteration had not as yet occurred 
within its small known range (4). 

CONSERVATION MEASURES Protected by law. Not known to occur in any reserve area and 
has never been studied (8). Surveys are needed to discover whether the species still survives 
and if so to recommend appropriate conservation measures; these will undoubtedly include 
protection of its habitat, and the possibilities of captive breeding. 



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An environmental profile of Madagascar 

All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on International 
Trade in Endangered Species of Wild Fauna and Flora, trade in them or their products is 
therefore subject to strict regulation and trade for primarily commercial purposes is banned. 

All Lemuroidae are listed in Class A of the African Convention, 1969, i.e., they may be 
hunted, killed, captured or collected only on the authorization of the highest competent 
authority, if required in the national interest or for scientific purposes. 

CAPTIVE BREEDING The species has never been kept in captivity. 

REMARKS For description of animal see (2,3,5,7,9). The species has been included in the 
family Lemuridae. 

REFERENCES 

1. Gunther, A. (1875). Notes on some mammals from Madagascar. Proceedings of the 
Zoological Society of London: 78-80. 

2. Napier, J.R. and Napier, P.H. (1967). A Handbook of Living Primates. Academic Press, 
London. 

3. Petter, A. and Petter, J. -J. (1971). Part 3.1 Infraorder Lemuriformes. In: Meester, J. and 
Setzer, H.W. (eds), The Mammals of Africa: An Identification Manual. Smithsonian 
Institution Press, City of Washington. 

4. Petter, J.-J. (1972). In litt. 

5. Petter, J.-J. and Petter-Rousseaux, A. (1956). A propos du lemurien 
malgache Cheirogaleus trichotis. Mammalia 20(1): 46-48. 

6. Petter, J.-J., Albignac, R. and Rumpler, Y. (1977). Mammiferes lemuriens (Primates 
prosimiens). Faune de Madagascar No. 44, ORSTOM-CNRS, Paris. 

7. Petter-Rousseaux, A. and Petter, J.-J. (1967). Contribution a la systematique des 
Cheirogaleinae (lemuriens malgaches). Allocebus, Gen. Nov., pour Cheirogaleus trichotis 
Gunther 1875. Mammalia 31(4): 574-582. 

8. Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy lemurs: Conservation or 
extinction? In: Tattersall, I. and Sussman, R.W. (eds), Lemur Biology. Plenum Press, New 
York. 

9. Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New York. 

10. Mittermeier, R.A. (1986). In litt. 



GREATER DWARF LEMUR 

Cheirogaleus major E. Geoff roy, 1812 Primates: Cheirogaleidae 

SUMMARY Endemic to Madagascar where it occurs in forested areas in the east. Not 
considered threatened at the present time although forest loss will undoubtedly decrease its 
range and numbers. Protected by law and occurs in a number of reserves. Has not been 
studied in the wild. 

DISTRIBUTION Madagascar. Occurs throughout the forested areas of the east from 
Taolanaro (Fort Dauphin) to Mt d'Ambre, and extending westward to include the Tsaratanana 
Massif and the Sambirano region (7). Petter el al. also indicate that a population of C. major 
exists on the Bongolava Massif, at the far western edge of the Eastern Region; presumably its 
isolation occurred relatively recently (3,7). Most recent authors have recognized at least two 
subspecies, of which one (C. m. crossleyi) is reported to occur primarily to the north of the 
Masoala Peninsula, while the other (C. m. major) is found to the south (3,7). However 
Tattersall believes the situation is still unclear and prefers at present to regard the species as 
variable but monotypic (7). The range of C. major once extended well onto the central 
plateau. For maps see (3,7). 



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Appendix 3: species accounts 

POPULATION Not thought to be threatened at the present time, although in 1975 Richard 
and Sussman reported it to be declining (along with almost all Malagasy .lemurs) (5). More 
information is required. 

HABITAT AND ECOLOGY The humid forests of the east. Nocturnal; rests during the day in 
constructed nests (7). Aestivates (6). Almost nothing is known of social organization; 
individuals are invariably sighted singly, although seem to sleep in groups. Has been reported 
in 'considerable' densities in certain areas of high food concentrations. Dwarf lemurs 
apparently feed on ripe fruit, nectar and pollen, with insects only occasionally taken; leaves are 
never eaten (3,7). 

THREATS Undoubtedly the loss of forest in the east will adversely affect this species but no 
specific reports of threats have been located. 

CONSERVATION MEASURES Protected by law. Known to occur in the Montagne d'Ambre 
National Park and in Natural Reserves 1 (Betampona), 3 (Zahamena), 4 (Tsaratanana), 1 1 
(Andohahela) and the Special Reserves of Nosy Mangabe and Perinet (5,8,9). Has not been 
studied and although this species is not as yet considered threatened, an ethoecology study 
would certainly be valuable. 

All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on International 
Trade in Endangered Species of Wild Fauna and Flora, trade in them or their products is 
therefore subject to strict regulation and trade for primarily commercial purposes is banned. 

All Lemuroidae are listed in Class A of the African Convention, 1969, i.e., they may be 
hunted, killed, captured or collected only on the authorization of the highest competent 
authority, if required in the national interest or for scientific purposes. 

CAPTIVE BREEDING In 1986 five individuals were held in captivity, at Duke University 
Primate Center in the U.S.A. (10). Local people in Taolanaro are known to have specimens as 
pets (8). 

REMARKS For description of animal see (1,2,3,7). 

REFERENCES 

1. Napier, J.R. and Napier, P.H. (1967). A Handbook of Living Primates. Academic Press, 
London. 

2. Petter, A. and Petter, J.-J. (1971). Part 3.1 Infraorder Lemuriformes. In: Meester, J. and 
Setzer, H.W. (eds), The Mammals of Africa: An Identification Manual. Smithsonian 
Institution Press, City of Washington. 

3. Petter, J.-J., Albignac, R. and Rumpler, Y. (1977). Mammiferes lemuriens (Primates 
prosimiens). Faune de Madagascar No. 44, ORSTOM-CNRS, Paris. 

4. Pollock, J.I. (1983). In litt. 

5. Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy lemurs: Conservation or 
extinction? In: Tattersall, I. and Sussman, R.W. (eds), Lemur Biology. Plenum Press, New 
York. 

6. Schilling, A. (1980). Seasonal variation in the fecal marking of Cheirogaleus medius in 
simulated climatic conditions. In: Charles-Dominique, P., Cooper, H.M., Hladik, A., 
Hladik, CM., Pages, E., Pariente, G.F., Petter-Rousseaux, A., Schilling, A. and 
Petter, J.-J. Nocturnal Malagasy primates: Ecology. Physiology and Behaviour. Academic 
Press, New York. 

7. Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New York. 

8. O'Connor, S. and Pidgeon, M. (1986). In litt. 

9. Mittermeier, R.A. (1986). In litt. 

10. Wright, P.C. (1986). In litt. 



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FAT-TAILED DWARF LEMUR 

Cheirogaleus medius E. Geoffroy,1812 Primates: Cheirogaleidae 

SUMMARY Endemic to Madagascar. Exhibits marked adaptations to survival in the dry, 
deciduous forests of the west and south. Local population densities can be very high, but, in 
common with other species of this region, it is likely to be affected by habitat destruction. 
Legally protected and occurs in several nature reserves. Its survival will depend on the 
protection of suitable areas of habitat. 

DISTRIBUTION Madagascar, where it is still found in most of the western and southern 
forests (8,17) from the Bay of Narinda to Taolanaro (17). The species may also have occurred 
in eastern and northern Madagascar and in the Sambirano region, in sympatry 
with Cheirogaleus major, but the present status of such populations is problematical (17). For 
map see (17). 

POPULATION No census has been undertaken; however the species is still widespread and is 
unlikely to be threatened at present. 

HABITAT AND ECOLOGY Inhabits the dry, deciduous forests of the west and south which 
have a 7-8 month dry season. The most characteristic feature of the species is its ability to 
'aestivate' for at least six consecutive months and, in some instances, up to eight months of the 
dry season. This appears to be an adaptive feeding strategy which consists of harvesting 
'surplus' food when available, and withdrawing into aestivation during the period of minimum 
production (4). The dormant period is spent inside deep holes of tree trunks where three to 
five individuals may be piled upon each other (4). In the Marosalaza Forest the species was 
observed to emerge at the end of November, just before the rainy season, and thus just as the 
period of intensive food production began (4). Since the existence of hiding places seems to be 
an essential condition of its survival, it tends to be associated with trees of a certain size 
(4,8). C. medius is chiefly an opportunistic frugivore, although insects are also an important 
component of the diet. Seasonal variation in diet follows food availability: nectars and fruits in 
November; fruits and an increasing proportion of insects in the period December - February. 
After this time, the proportion of fruits in the diet may increase at the time of fattening 
preceding dormancy (4). Nocturnal. Individuals are invariably sighted singly (17). Petter has 
noted extremely high population densities of the species in the Marosalaza Forest, i.e. 350 
animals per 100 ha (4). Home ranges were about 4 ha. Mating was observed in the wild at the 
beginning of November and births occurred in January (4). Captive studies show gestation to 
be 61-64 days and litter size to be usually two offspring (range 1-4); cannibalism of newborn 
was frequent (2). Infants reach sexual maturity during their first year i.e. they are able to 
breed in their first breeding season (1). The genus Cheirogaleus is the only primate which 
regularly uses faeces as scent marking material (15). 

THREATS In 1972 Petter reported that forest and scrub clearance were causing a rapid 
contraction of range (8). 

CONSERVATION MEASURES Protected by law, although difficult to enforce. Known to 
occurs in Natural Reserves 7 (Ankarafantsika) and 11 (Andohahela) (9,14), as well as in the 
private reserves of Analabe and Berenty owned by M. de Heaulme (7,11,14,17). Has been 
studied to some extent in both the wild and captivity (1,2,3,4,15). This species is reputed to be 
particularly dependent on trees with holes in which it can hibernate, and this tends to be 
associated with trees of a certain age; habitat protection and management can thus be expected 
to be especially important. 

All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on International 
Trade in Endangered Species of Wild Fauna and Flora, trade in them or their products is 
therefore subject to strict regulation and trade for primarily commercial purposes is banned. 

All Lemuroidae are listed in Class A of the African Convention, 1969, i.e., they may be 
hunted, killed, captured or collected only on the authorization of the highest competent 
authority, if required in the national interest or for scientific purposes. 

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Appendix 3: species accounts 

CAPTIVE BREEDING In 1986 there were over 100 individuals in captivity, most at the Duke 
University Primate Center, U.S.A. (16). The majority are evidently captive bred (6). 

REMARKS For description of animal see (5,6,7,10,12,17). The species is often divided into 
two subspecies but according to Tattersall the distinction is unwarranted (17). 

REFERENCES 

1. Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, CM., Pages, E., Pariente, G.F., 
Petter-Rousseaux, A., Schilling, A. and Petter, J. -J. (1980). Nocturnal Malagasy 
Primates: Ecology, Physiology and Behaviour. Academic Press, New York. 

2. Foerg, R. (1982). Reproduction in Cheirogaleus medius. Folia Primatologica 39(1-2): 
49-62. 

3. Foerg, R. and Hoffmann, R. (1982). Seasonal and daily activity changes in 
captive Cheirogaleus medius. Folia Primatologica 38(3-4): 259-268. 

4. Hladik, CM., Charles-Dominique, P. and Petter, J. -J. (1980). Feeding strategies of five 
nocturnal prosimians in the dry forest of the west coast of Madagascar. In: 
Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, CM., Pages, E., Pariente, G.F., 
Petter-Rousseaux, A., Schilling, A. and Petter, J. -J. Nocturnal Malagasy Primates: 
Ecology, Physiology and Behaviour. Academic Press, New York. 

5. Napier, J.R. and Napier, P.H. (1967). A Handbook of Living Primates. Academic Press, 
London. 

6. Olney, P.J.S. (ed.) (1986). International Zoo Yearbook 24/25. Zoological Society of 
London. 

7. Petter, A. and Petter, J. -J. (1971). Part 3.1 Infraorder Lemuriformes. In: Meester, J. and 
Setzer, H.W. (eds), The Mammals of Africa: An Identification Manual. Smithsonian 
Institution Press, City of Washington. 

8. Petter, J.-J. (1972). In lilt. 

9. Petter, J.-J. and Petter-Rousseaux, A. (1956). A propos du lemurien 
malgache Cheirogaleus trichotis. Mammalia 20(1): 46-48. 

10. Petter, J.-J., Albignac, R. and Rumpler, Y. (1977). Mammiferes lemuriens (Primates 
prosimiens). Faune de Madagascar No. 44, ORSTOM-CNRS, Paris. 

11. Petter, J.-J., Schilling, A. and Pariente, G. (1971). Observations eco-ethologiques sur 
deux lemuriens malgaches nocturnes: Phaner furcifer et Microcebus coquereli. La Terre el 
la Vie 118(3): 287-327. 

12. Petter-Rousseaux, A. and Petter, J.-J. (1967). Contribution a la systematique 
des Cheirogaleinae (lemuriens malgaches). Allocebus, Gen. Nov., pour Cheirogaleus 
trichotis Gunther 1875. Mammalia 31(4): 574-582. 

13. Richard, A. (1983). In lilt. 

14. Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy lemurs: Conservation or 
extinction? In: Tattersall, I. and Sussman, R.W. (eds), Lemur Biology. Plenum Press, New 
York. 

15. Schilling, A. (1980). Seasonal variation in the fecal marking of Cheirogaleus medius in 
simulated climatic conditions. In: Charles-Dominique, P., Cooper, H.M., Hladik, A., 
Hladik, CM., Pages, E., Pariente, G.F., Petter-Rousseaux, A., Schilling, A. and Petter, 
J.-J. Nocturnal Malagasy Primates: Ecology. Physiology and Behaviour. Academic Press, 
New York. 

16. Wright, P.C. (1986). In lilt. 

17. Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New York. 

18. O'Connor, S. and Pidgeon, M. (1986). In lift. 

19. Durrell, L. (1986). Pers. comm. 



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An environmental profile of Madagascar 
COQUEREL'S MOUSE LEMUR 

Mirza coquereli (A. Grandidier, 1867) Primates: Cheirogaleidae 

SUMMARY Endemic to the forests of western Madagascar which are fast disappearing. 
Protected by law, has been the subject of several etho-ecology projects, and occurs in two 
protected areas. A captive breeding group exists. 

DISTRIBUTION Madagascar where it has a restricted range in the forests of the west 
(5,8,10,13), apparently in a scattered series of disjunct isolates (13), principally between the 
Onilahy and Fierenana Rivers, i.e. from about the region of Ankazoabo northwards to 
Belo-sur-Tsiribihina, or a little beyond (13). It also occurs on the Ampasindava Peninsula and 
in the adjoining region (13). Petter el al. have suggested that it occurs in coastal forests in the 
area intervening between the southern and northern populations (9), but this remains 
unconfirmed (13). Although the former range is not precisely known it was certainly more 
extensive than today (8). For maps see (9,13). 

POPULATION Mittermeier (1986) commented that it was probably endangered; its range is 
very small and forest loss is rapid (14). In 1975 Sussman and Richard considered M. coquereli 
to be one of the Malagasy lemurs which was 'extremely rare and probably on the brink of 
extinction' (12). In 1972 it was reported to be still fairly abundant in some isolated forest 
patches (8), for example along the streams of the coastal forest north of Morondava, although 
was considered threatened (8). 

HABITAT AND ECOLOGY Inhabits the more humid parts of the dry, deciduous forests of 
western Madagascar. Adapts to the dry season, when forest productivity is markedly 
diminished, by feeding mainly on the secretions produced by colonies of homopteran larvae; 
during the wet season feeds omnivorously: on fruit, flowers, and insects, and probably also on 
eggs and small vertebrates (1,5,13). Nocturnal; during the day rests in large spherical nests 
which it constructs of small twigs (5,8), usually located at a height of 2-10 m, often in trees 
which do not shed their leaves during the dry season (e.g., Euphorbiaceae) and are thickly 
covered with lianas (5). Pages reports that contact between individuals during the night's 
activity is rare, although she suggests that a 'loose pair bonding' between males and females 
may exist. Male and female ranges overlap partially, that of the female averages about 10 ha, 
that of the male about 8 ha. Most time is, however, spent in much smaller core areas, 
averaging 3 ha for females and 2 ha for males. Pages believes that the core area is defended, at 
least by males (4,5,6,13). The species exhibits only a brief period of sexual activity, gestation 
lasts about three months. It is reported to have 2-3 young a year (8), though animals at Duke 
University have a litter size of 1 or 2 (11). 

THREATS The species's range is contracting through forest degradation and destruction, and 
by cultivation of the remaining areas of land that can be irrigated (8). 

CONSERVATION MEASURES Legally protected but enforcement is difficult. It is known to 

occur in Natural Reserve 9 (Tsingy de Bemaraha) and in Analabe private reserve north of 

Morondava owned by M. de Heaulme (12). The species has been the subject of a number of 
studies (4,5,6,10). 

All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on International 
Trade in Endangered Species of Wild Fauna and Flora, trade in them between acceding nations 
is therefore subject to strict regulation and trade for primarily commercial purposes is banned. 

All Lemuroidae are listed in Class A of the African Convention, 1969, i.e., they may be 
hunted, killed, captured or collected only on the authorization of the highest competent 
authority, if required in the national interest or for scientific purposes. 

CAPTIVE BREEDING One breeding colony exists, at Duke University Primate Center; in 
September 1986 it comprised 23 animals (2). In the early 1970s a pair were also kept for 
breeding purposes at the Brunoy Laboratory of the French Museum of Natural History (8). 



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Appendix 3: species accounts 

REMARKS For description of animal see (3,7,9,13). Often included in the genus Microcebus 
(e.g. 4,5). 

REFERENCES 

1. Hladik, CM., Charles-Dominique, P. and Petter, J. -J. (1980). Feeding strategies of five 
nocturnal prosimians in the dry forest of the west coast of Madagascar. In: 
Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, CM., Pages E., Pariente, G.F., 
Petter-Rousseaux, A., Schilling, A. and Petter, J.J. Nocturnal Malagasy Primates: Ecology, 
Physiology and Behaviour. Academic Press, New York. 

2. Wright, P.C (1986). In litt. 

3. Napier, J.R. and Napier, P.H. (1967). A Handbook of Living Primates. Academic Press, 
London. 

4. Pages, E. (1978). Home range, behaviour, and tactile communication in a nocturnal 
Malagasy lemur, Microcebus coquereli. In Chivers, D.J. and Joysey, K.A. (eds), Recent 
Advances in Primatology. Vol. 3. Academic Press, New York. 

5. Pages, E. (1980). Ethoecology of Microcebus coquereli during the dry season. In: 
Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, CM., Pages, E., Pariente, G.F., 
Petter-Rousseaux, A., Schilling, A. and Petter, J. -J. Nocturnal Malagasy Primates: 
Ecology, Physiology and Behaviour. Academic Press, New York. 

6. Pages, E. (1980). Relations sociales et importance des echanges tactiles dans la cohesion 
familiale chez Microcebus coquereli, lemurien malgache primitif. Acad. Malgache. 

7. Petter, A. and Petter, J. -J. (1971). Part 3.1 Infraorder Lemuriformes. In: Meester, J. and 
Setzer, H.W. (eds), The Mammals of Africa: An Identification Manual. Smithsonian 
Institution Press, City of Washington. 

8. Petter, J.-J. (1972). In litt. 

9. Petter, J.-J., Albignac, R. and Rumpler, Y. (1977). Mammiferes lemuriens (Primates 
prosimiens). Faune de Madagascar No. 44, ORSTOM-CNRS, Paris. 

10. Petter, J.-J., Schilling, A. and Pariente, G. (1971). Observations eco-ethologiques sur 
deux lemuriens malgaches nocturnes: Phaner furcifer et Microcebus coquereli. La Terre et 
la Vie 118(3): 287-327. 

11. Pollock, J.I. (1983). In litt. 

12. Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy lemurs: Conservation or 
extinction? In: Tattersall, I. and Sussman, R.W. (eds), Lemur Biology. Plenum Press, New 
York. 

13. Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New York. 

14. Mittermeier, R.A. (1986). In litt. 



GREY MOUSE LEMUR 

Microcebus murinus (J.F. Miller, 1777) Primates: Cheirogaleidae 

SUMMARY Endemic to Madagascar. This and its sister taxon Microcebus rufus are the most 
abundant of the lemurs and are not considered threatened at the present time. Protected by 
law, has been the subject of studies, and occurs in a number of reserves. 

DISTRIBUTION Madagascar. Occurs throughout the forested areas of western, southern and 
south-eastern Madagascar, from Taolanaro to the Sambirano region (15). Martin (1972) notes 
that in the Taolanaro area the Grey Mouse Lemur's area of distribution includes the littoral 
forest to the north and east of the town, while the Brown Mouse Lemur Microcebus rufus, 
occurs in the rain forest which extends southward, to the west of Taolanaro, almost to the 
coast. There is thus in this region a sharp environmental demarcation between the two species, 
since although the two areas receive similar rainfall, the littoral is much better drained, and 
supports a vegetation of distinctly less humid aspect that does the interior (6,15). The northern 
boundary of the Grey Mouse Lemur is not precisely known, but the species appears not to 
occur north of the Sambirano River (15). Grey Mouse Lemurs are also found on Nosy Be 
Island off the north-west coast (3). For maps see (12,15). 

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An environmental profile of Madagascar 

POPULATION Not threatened; the species remains widespread and abundant. In 1975 M. 
murium and M. rufus were considered (together) to be the most abundant lemurs and perhaps 
the only ones not declining in number (14). However, even for M. murinus Martin (1973) gives 
data indicating that heavy tree-felling had been affecting population densities (7,8). A study 
in the south also suggested that population densities were not as high as previously estimated 
and that extensive grazing by both cattle and goats was destroying the low bush habitat of the 
species even in areas where no actual tree felling occurred (8). 

HABITAT AND ECOLOGY Forests of western Madagascar, where it seems to be adapted to 
forest-edge habitats. Martin has noted that Microcebus are usually sighted within a few metres 
of paths and tracks and rarely deep in the forest (6,7,15). In arid Didierea forest near 
Bemarivo, not far from Mandena, the species occurred at a density of 3.6 individuals per ha 
(2,15). Nocturnal; they sleep in nests during the day. Seemingly solitary when active but sleep 
in groups. Martin has suggested that the species forms relativ